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Memoirs Of the Number 37

NATIONAL MUSEUM

Of Victoria Melbourne Australia 30 June 1976
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Cover — An Obulkarra woman being painted with a clan
design after death.

MEMOIRS

of the

NATIONAL MUSEUM OF VICTORIA

MELBOURNE AUSTRALIA

No. 37

Director
J. McNally

Deputy Director
Thomas A. Darragh

Editor
Douglas M. Stone

PUBLISHED BY ORDER OF THE TRUSTEES
30 JUNE 1976

Brown Prior Anderson Pty Ltd 5 Evans Street Burwood 3125

NATIONAL MUSEUM OF VICTORIA

Members of Council

Sir Robert Blackwood, MCE BEE FIE Aust (President)
Henry G. A. Osborne, BAgrSc (Deputy President)
James C. F. Wharton, BSc (Treasurer)

Professor E. S. Hills, CBE PhD (Lond) Hon DSc (Dunelm) DSc FIC FAA

FRS
Professor Sir Sydney Sunderland, CMG MD BS DSc FRACP FRACS FAA
Dr. H. K. Worner, DSc (Melb) Hon DSc (Newcastle) ABSM FAA FRACI

FIM F1MM MA1MM MAIME FTS
Sir Henry Somerset, CBE MSc FRACI MAIMM
Brigadier P. P. Jackson, CMG CBE BE FIE (Aust) FI Mech E FAIM
Professor J. W. Warren, MA PhD (Calif)

John McNally, ED MSc, Chief Executive Officer
W. L. Drew, Secretary to Council

Staff (as at 23/2/1976)

Director: John McNally, ED MSc AMAA

Deputy Director: Thomas A. Darragh, MSc DipEd AMAA

Administration:

A. G. Parsons (in charge)

E. J. Peat

E. Rowley

Patricia Batchelor

Enid Howell

Kayelene Voege

Lynette Anderson

Scientific Staff

Geology and Palaeontology:

Curator of Fossils: T. H. Rich, PhD BA

Curator of Minerals: W. D. Birch, BSc (Hons)

Assistant Curator of Fossils: Vacant

Assistants: F. Bussat, I. Stewart, FRMIT (Geol), Susan Gibson

Vertebrate Zoology:

Curator of Vertebrates: Joan M. Dixon, BSc (Hons)
Curator of Birds: A. R. McEvey, BA
Assistants: Linda Chegwidden, BSc, I. T. Beck

Herpetology: A. J. Coventry

Invertebrate Zoology: ^™

Curator of Insects: A. Neboiss, MSc FRES
Assistant Curator of Insects: Vacant
Curator of Invertebrates: B. J. Smith, PhD BSc AMAA
Assistant Curator of Invertebrates: Suzanne Stevenson BSc
Assistants: Elizabeth Matheson, Rhyllis J. Plant

Antk Cuvltov y: oi Anthropology: A. L. West, BA DipSocStud
Assistant Curator: Alison M. Oates, BA
Assistants: D. S. Wood, Christine Hogarth, BA

Survey Officer:

J. D. Blyth, BSc

Assistants: R. Houghton, Jane Avery

Library: .

Clara Myers, BA, DipLib

Lorna Crowther, BSc, DipLib

Assistants: Josie Pearce, Leeanne ODonnell

Display and Preparation:

P C. Duce, DipAD, Display Officer
L. J. Chapman, Senior Preparator
M. G. Traynor
B. Hall
K. Kelly

S. Huxley, Dip AD

Diane Stephens, DipArt

Jan Wheeler ASTC (DipGraphDesign)

Photographer:

F. Coffa

Education Service:

J. Woollard, TPTC (in charge)
Brenda Traynor, TITC
R. L. Miller, DipArt

G. Wallis, FRMIT (Geol) TSTC

D. Poad, BA DipEd
Rosemary Taylor, BA TPTC
Linda Dunn, TPTC
Valerie McCormack, (typist)

Honorary Associates

with year of appointment

Geology:

A. A. Baker, 1951

A. W. Beasley, MSc PhD DIC, 1973

A. C. Collins, FRAIA ARIBA AMTPI MACE, 1953

E. D. Gill, ISM BA BD FGS FRGS, 1973
Professor J. F. Lovering, MSc PhD, 1974
J. A. Talent, MSc PhD, 1966

D. J. Taylor, MSc, 1966
H. E. Wilkinson, BSc, 1970

Vertebrate Zoology:

C. N. Austin, 1955

C. W. Brazenor, 1962

A. G. Brown, MRCS (Eng) LRCP (London), 1968

R. P. Cooper, FNIA, 1952

N. J. Favaloro, 1945

A. K. Lee, BSc (WA) MA PhD (Calif), 1972

M. J. Littlejohn, PhD (WA) MSc, 1972

P. A. Rawlinson, BSc, 1968

C. Tanner, 1963

R. M. Warneke, MSc, BAgrSc, 1966

H. N. B. Wettenhall, MD BS MRCP FRACP, 1963

Invertebrate Zoology:

K. N. Bell, BSc DipEd, 1973
J. Hope Black, MSc, 1966
R. F. Burn, 1962
A. N. Burns, MSc FRES, 1966

D. F. Crosby, FRES AASA AFAIM, 1968
R. L. Jensz, BSc DipEd, 1968

C. McCubbin, 1974

A. E. Monger, LS MIS (Aust), 1974

E. T. Smith, 1960

Jeanette E. Watson, ASMB AMTC, 1970

L. Winsor, DIP MED TECH FAIMT AAIST, 1974

Anthropology:

D. A. Casey, MC FSA, 1933

J. H. McNamarra, MB BS FRCPA, 1969
N. M. Wallace, 1970

CONTENTS

PAPERS

PALAEONTOLOGY

1. A revision of the Gastropod Fauna of the Lily dale Limestone (Early
Devonian) of Victoria. By C. B. Tassell. (Plates 1-3) 1

VERTEBRATE ZOOLOGY

2. A new species of Hemiergis (Scincidae: Lygosominae) from Victoria. By

A. J. Coventry. 23

3. The Endemic Australian Lizard Genus Morethia (Scincidae: Lygoso-
minae) in southern Australia. By P. A. Rawlinson. 27

INVERTEBRATE ZOOLOGY

4. A creeping Ctenophoran (Platyctenea: Ctenophora) from Victoria, Aus-
tralia. By Brian J. Smith and Rhyllis J. Plant. 43

5. Symbiocladius aurifodinae sp. nov. (Diptera, Chironomidae), A Parasite
of Nymphs of Australian Leptophlebiidae (Ephemeroptera). By H. B. N.
Hynes. 47

6. The Ascidian Fauna of Western Port, Victoria and a comparison with that

of Port Phillip Bay. By Patricia Kott. 53

ANTHROPOLOGY

7. Mortuary Customs of northeast Arnhem Land: an account compiled from
Donald Thomson's fieldnotes. By Nicolas Peterson. (Plates 4-11) 97

A REVISION OF THE GASTROPOD FAUNA OF THE LILYDALE
LIMESTONE (EARLY DEVONIAN) OF VICTORIA

By C B. Tassell

* Present Address Albany Residency Museum, Port Road, Albany, W. A.
Assistant Curator of Fossils, National Museum of Victoria

Abstract

Thirteen species of gatsropods are described from the Lower Devonian Lilydale Limestone
near Melbourne. Two of these are type species of the genera Scafaetrochus (S. lindstrotni
Etheridge) and Gyrodoma (G. etheridgei (Creswell)). Another two species Michelia brazieri
(Etheridge) and in part Straparollus {Euomphalus) northi (Etheridge) are the type species
for two now synonymized genera Vetotuba and Liomphalus. The other species are Tremanotus
pritchardi Cresswell, Bellerophon (B.) cresswell Etheridge, Phanerotrema australis Etheridge,
Stenoloron subaequilatera (Chapman), Naticopsis (N.) lilydalensis Cresswell, Murchisonia
(M.) pritchardi (Etheridge), Siluriphorus antiquus (Cresswell), Loxonema australis (Chap-
man), and Oriostoma rotundimuratus sp. nov. Also described is Michelia danvhii (de Koninck)
from the Lower Devonian 'Receptaculites' Limestone, Taemas, New South Wales. The genus
Boiotremus Horny is considered to be a synonym of Tremanotus Hall.

The fauna which lacks platyceratids is associated with a diverse coral and stromatoporoid
assemblage and depleted brachiopod fauna. The gastropod fauna possesses strong affinities
with the Old World Realm faunas of Europe and North America, and also indicates a
continuation into the Lower Devonian of certain typically Silurian forms.

Introduction

The gastropod fauna of the Lilydale Lime-
stone was the subject of much attention by
early palaeontologists in Australia. Robert
Etheridge, Junr. ? in a series of papers (1890,
1891, 1894 and 1898) described three new
genera, Gyrodoma, Scalaetrochus and Veto-
tuba, nine new species and noted the presence
of an operculum in place in Straparollus
{Euomphalus) northi. At about the same time
the Rev. A. W. Cresswell (1885, 1893 and
1894), while noting the general elements of
the fauna described four new species and noted
the presence of an operculum in place in 5.
(E.) northi.

Soon after, Chapman (1916) reviewed the
entire gastropod fauna. He described one new
genus Liomphalus, six new species or varieties,
and noted the presence of Omphalotrochus
globosum (Schlotheim). Subsequently the fauna
received very little attention. Knight (1941)
reviewed the genera established by Etheridge
and Chapman; Philip and Talent (1959) dis-
cussed S. (E.) northi and Scalaetrochus lind-
stromi and more recently Yochelson and Lin-
sley (1972) discussed the opercula of S. (E.)
northi and Cyclonema lilydalensis.

The gastropod fauna of the limestones in the

Taemas region, N.S.W., has had a similar
history of early attention and subsequent neg-
lect. De Koninck (1876) described one new
genus, Mitchellia, which Knight et al. (p. 1301,
1960) synonymized with Scoliostoma, seven
new species and noted the presence of six
previously described species.

The basis of this study was the large col-
lection of the National Museum of Victoria
which has been collected over a period of
more than seventy years. The bulk of this has
come from the limestone where it has been
the subject of Tertiary weathering (O. P.
Singleton pers. comm.). Fortunately the wea-
thering process preferentially destroys the
matrix of the limestone before destroying the
fossils. Thus at the right stage in this process
when the matrix is soft the fossils can be
obtained free of matrix and moderately well
preserved. The remainder of the collection is
preserved in a dense grey limestone.

Middle Palaeozoic gastropods have been the
subject of relatively little recent study both in
Australia and overseas. As a consequence many
of the genera have been interpreted in a very
loose sense. So much so, that quite frequently
authors note that the species they are discussing,
while being assigned to an established genus

C. B. TASSELL

in fact belongs to a new genus. In view of this,
each of the species from Lilydale has been
compared with the type species of the appro-
priate genus.

However, this has not been done with Cyclo-
nema australis Etheridge and C. lilydalensis
Etheridgc, the two species of this genus de-
scribed from Lilydale. It is generally recognized
that the Devonian forms assigned to the genus
Cyclonema in fact constitute at least one new
genus (Thompson, 1970). To date no one has
attempted to erect a new genus for these forms
because a satisfactory basis for distinguishing
it from Cyclonema sensu stricto is not readily
apparent. However, the palaeoecological impli-
cations arising from the assignment of these
species to the genus Cyclonema are quite
erroneous. Thus the species from Lilydale are
not redescribed here, rather it is intended that
they should be the subject of another study.
Also not described are members of the classes
Monoplacophora, Polyplacophora, Pelecypoda,
and Rostroconchia known to occur at Lilydale.
These are to be the subject of another paper.

The generic ranges and distributions given
by Knight et al. ( 1960) for the genera discussed
here are accepted in general. Any amendment
of this is based upon a sensu-sJricto interpreta-
tion of the genus.

In this study the following abbreviations
have been used: P: Palaeontological collection
of the National Museum of Victoria; F.:
Australian Museum, Sydney; M.U.G.D.:
Melbourne University Geology Department;
A.N.U.: Geology Department, Australian Na-
tional University.

All measurements are in millimetres and the
following symbols relating to the measurements
have beein used:
Clu, spiral sculptural elements above the seleni-

zone.

Gil, spiral sculptural elements below the selcni-

zonc.
Hap, height of aperture.
Ht, total height of shell.
L, length measured at the sclenizone in bellero-

phontids.
Lap, length of aperture.
Sw, selenizonc width.

Wap, width of aperture.

Wit, width at last trema.

Wt, total width of shell.

Wh, total number of whorls in shell.

*, specimen incomplete.

Acknowledgements

I wish to thank Dr A. Ritchie of the Austra-
lian Museum, Sydney, who made available
Etheridge's type material; Mr M. Cooper of
Melbourne University Geology Department,
for the loan of specimens; Mr T. A. Darragh
and Dr K. S. W. Campbell for critically reading
this manuscript and their helpful comments and
discussion; Dr O. P. Singleton for his discussion
and comments throughout the course of this
study; Margaret Tassell for her discussions and
encouragement and Mr F. Coffa for the photo-
graphs.

Age of the Faunas

In the past, the age of the Lilydale Lime-
stone has been the subject of some contention
and frequent revision. Strusz (1972) assessed
the evidence presently available from a number
of groups and regarded the Lilydale Limestone
as being Late Sicgenian in age.

Strusz (1972) considered the 'Receptaculites'
Limestone at Taemas, in which Michelia dar-
wini (de Koninck) occurs, to be Emsian in age.

However, Philip (1974) summarized recent
developments in Europe which have placed the
relationship of the stages in the different facies
of the Lower Devonian in a state of flux. He
commented, 'How sterile now seems the de-
bates as to whether certain limestone horizons
in eastern Australia are Siegenian or Emsian
(or even Pragian) in age'. As yet the relation-
ship of these stages in Europe has not been
resolved.

This revision of the gastropod fauna makes
little contribution to the age determination
because of the present inadequate knowledge
of gastropod faunas both in Australia and
overseas. Most of the genera represented at
Lilydale are characterized by such long ranges
that they are of little value in age determina-
tions. The remaining few genera with relatively
short ranges such as Scalaetrochus are of limited
value because of their restricted distribution.

REVISION OF THE GASTROPOD FAUNA

Relationships of the Fauna

Representatives of eight gastropod super-
families, murchisoniaceans, euomphalaceans,
bellerophontaceans, pseudophoraceans, pleuro-
tomariaceans, oriostomataceans, neritaceans
and loxonemataceans are described from the
limestone at Lilydale. As well as these there
are the two species of turbiiniform gastropods
previously assigned to the genus Cyclonema.

Comparison with other Lower Devonian
faunas in Australia is limited by the notable
lack of recent studies. The closest fauna geo-
graphically that is described, is that of the
Marble Creek Limestone (Talent and Philip,
1956). This fauna is dominated by platycera-
tids, although species of Tremanotus and
Michelia are known at Marble Creek as well as
Lilydale. Chapman (1907, p. 73) also noted
the presence of Scalaetrochus sp., but this has
not been subsequently verified.

The gastropod fauna from the limestones at
Taemas, N.S.W., as described by de Koninck,
has only a limited number of genera in com-
mon with Lilydale. Michelia darwini, described
here, is very similar in shape to M. brazieri
from Lilydale. Although de Konimck only de-
scribed small bellerophontids from Taemas such
as Bellerophon convolutus de Koninck, large
forms comparable in size to B. (B.)cresswelli
Etheridge are known to occur as well. Other-
wise the gastropod faunas differ considerably,
with Taemas possessing an abundance of small
high spired forms, whereas such forms are
generally absent at Lilydale.

Boucot (1975) summarized the now con-
siderable amount of work on Devonian palaeo-
geography. During t he Lower and Middle
Devonian a very marked provincialism de-
veloped when compared with the preceding
Silurian period and succeeding Late Devonian.
This provincialism was greatest during the
Siegenian, Emsian and Eifelian. During the
Siegenian three realms, Malvinokaffric, Old
World and Eastern Americas have been recog-
nized, principally on the basis of brachiopods.
Other groups including trilobites, corals and
conodonts also support this biogeographic
scheme. Within the Old World realm a number

of regions or sub-provinces are recognized. It
is within one of these, the Tasman sub-pro-
vince, that Lilydale is located.

Boucot has observed that during 'the early
Devonian the oriostomatids — poleumitids, tre-
manotids and euomphalids are present only
in the Old World Realm as "Silurian holdovers"
(relicts).' They also tend to be particularly
characteristic of the Bohemian facies of the
Old World Realm.

The gastropod fauna at Lilydale strongly
supports this palaeobiogeographical scheme.
Fifteen per cent of the fauna consists of relict
genera Tremanotus, Straparollus (Euomphalus)
and Oriostoma. These relict genera also tend
to emphasize the greater similarity between
Lilydale and the gastropods of the Bohemian
facies, rather than with the sandier Rhenish
facies of the Rhenish-Bohemian sub-province.
The dextrally coiled S. (E.) northi is very like
the similarly coiled S. (E.) carnicus (Freeh)
from the Carnac Alps. Also present in the Low-
er Devonian limestones of the Carnic Alps are
species of Bellerophon, Phanerotrema and
Stenoloron which are very similar to the species
of these genera at Lilydale. Spitz (1907, pi. 13,
fig. 16 a and b) illustrated a species of Poly-
tropis from the Carnic Alps which is closely
similar to the species assigned to 'Cyclonema*
at Lilydale. Jhaveri (1969) considered that
the Carnic Alps gastropod fauna shows a close
relationship with the gastropod faunas from
the Lower Devonian of Bohemia, Northern
France and New York as does the fauna
from Lilydale.

Palaeoecology

Two gastropod groups dominate the fauna,
the murchisoniaceans and the species of 'Cyclo-
nema'. They constitute about 31% a:nd 25%
of the fauna respectively. The euomphalaceans,
bellerophontaceans and pseudophoraceans are
the next most abundant groups in the fauna
comprising 13%, 11% and 9% respectively.
The remaining four superfamilies each com-
prise about 3% of the fauna.

In terms of general shape, the turbiniform
to high spired forms comprise about 70% of

C. B. TASSELL

the fauna and the other 30% consists largely of
discoidal and planispiral forms. About 80% of
the fauna is cither medium sized or large,
although this may in part be an artifact of
selective collecting. However, field comparisons
of the abundance of smaller forms at Lilydale,
Buchan and Taemas indicate a very much
greater abundance at the latter two localities
than at Lilydale.

At Lilydale the most distinctive feature of
the fauna is the total absence of members of
the Family Platyccratidae which is presently
characterized by a cophrophagous mode of
life. The two species of 'Cyclonema* described
from Lilydale possess an operculum (Yochel-
son and Linsley, 1972), lack any apertural or
coiling irregularity and possess no other feature
that could be suggestive of a cophrophagous
mode of life. Thus their association with the
platyccratidae would greatly distort any inter-
pretation of the environment of the fauna.

Not only is there a general absence of platy-
ceratids at Lilydale, there is also a general lack
of crinoidal remains in the limestone. A signifi-
cant percentage of the crinoidol remains present
arc composed of fragments of Pcmerocrinus
(Bates, 1972). This is in marked contrast to
the gastropod fauna of the Marble Creek lime-
stone. This latter fauna, dominated by platy-
ccratids, occurs in a limestone composed in
large part of crinoidal fragments. The fauna is
also notable for its low taxomomic diversity in
comparison with the diversity at Lilydale.

A similar situation is seen in the diversity of
the gastropod fauna in the Silurian recfal com-
plexes of Gotland, Sweden. The reefal lime-
stones in which Manten (1971) observed
gastropod faunas had a considerably more
diverse fauna than that of the associated cri-
noidal limestones.

Linsley (1968) in his description of the
Middle Devonian gastropods of the Anderdon
Limestone recognized two principal habitats.
One of these is a 'biostromal' environment
where small snails lived on the carbonate mud
flats between the corals and stromatoporoids in
relatively shallow water. The gastropods of the
other habitat, that of the 'inter-reef lived on
the carbonate muds developed between sparse,

localized tetracoral assemblages. This fauna
was distinguished by its considerably larger
size, l"-5" being the maximum dimension. Also
present were a few nautiloids, articulate
brachiopods and some ostracodes. Linsley
considered that the fauna and the sedimeait
suggested a fairly quiet environment.

The composition of the fauna of Lilydale
resembles reasonably closely the fauna of Les-
ley's inter-reef habitat. Both are dominated by
large forms although more of these are high-
spired types at Lilydale. However, the first spec-
ies of Scalaeirochus to be recorded outside
south-eastern Australia was described by Lins-
ley from the Anderdon Limestone. Thus the
similarity of sediment and fauna suggest that
the gastropods of the Lilydale Limestone oc-
cupied an environment somewhat similar to
the inter-reef environment of the Anderdon
Limestone.

Yochelson and Dutro (1960) in their de-
scription of a Mississippian and Permian
gastropod fauna from limestones in northern
Alaska also made some comments on the
palaeoecology of the assemblages. They ob-
served that 'Platyceras commonly occurs here
in crinoidal limestones'. This is in accord-
ance with its distribution in the limestones at
Gotland, Marble Creek and Lilydale. Where
Platyceras is common m the Lower Mississip-
pian sediments, 'the associated gastropods show
less variety than in the Upper Mississippian'
(where Platyceras is less common). Again this
is comparable with the situation at Lilydale
and Gotland.

However, they noted that 'corals and gastro-
pods also appear to be nearly mutually exclu-
sive' and that 'gastropods are commonly as-
sociated with numerous taxonomically diver-
sified brachiopods'. At Lilydale a situation
quite the reverse exists. Gastropods and corals
are very closely associated while only a de-
pleted and restricted brachiopod fauna is pre-
sent.

That such diverse gastropod associations
exist only serves to emphasize the need for
further work on this group before more mean-
ingful generalizations on gastropod palaeoe-
cology can be made.

REVISION OF THE GASTROPOD FAUNA

Systematic Descriptions

Superfamily BELLEROPHONTACEA

McCoy, 1851
Family Sinuitidae Dall in Zittel-Eastman,

1913
Subfamily Tremanotinae Peel, 1972
Genus Tremanotus Hall, 1865

( = Boiotremus Horny, 1962).

Type Species; Tremanotus alpheus Hall, 1865;
Middle Silurian; Bridgeport, Illinois, U.S.A.
Range: Middle Ordovician to Lower Devonian.
The presence of 7\ pritchardi at Lilydale and
T. fortis Freeh and T. insectus Freeh in the
Upper Koneprusy Limestone, Koneprusy ex-
tends the upper range of the genus from Middle
Silurian to Lower Devonian.
Discussion: Knight et al. (1960) provided a
diagnosis for this genus in which the slit is
'represented by a row of tremata, all but the
last few closed, not extending on to expanded
lip'. Subsequently Horny (1963) amended this
diagnosis to read, 'slit represented by a row of
tremata in body whorls, not extending on the
expanded lip; no tremata but shallow sinus in
outer lip in young stages'. Horny (1962) also
erected a new genus, Boiotremus, characterized
by 'tremata present along the whole length of
the whorls, periodical widened apertures after
distances of 1-3 tremata'. It is into this latter
genus that most of the species previously as-
signed to Tremanotus would be placed.

In his discussion of the genus Tremanotus
he noted that the 'main and characteristic sign
is the existence of the five opened tremata in
the body whorl region. In the ontogenetically
younger stage there are no tremata developed
, . .' This interpretation is inconsistent with T.
alpheus as described by Hall and Knight's
(1941) redescription which Horny (1963, p.
97) suggested implicitly supported his case.
Knight (noted '6-8 tremata remaining open' and
'the earlier ones (were) filled'. That there are
more than 5 tremata can be clearly seen in
Knight's figures of the type species. The figures
of Clarke and Ruedemann (1903) also clearly
indicate that the presence of tremata is not
confined to the body whorl. Unfortunately the
preservation and orientation of the specimens

figured by Horny does not show the dorsal
surfaces of the earlier whorls.

Thus Horny's amendment of the diagnosis
of Tremanotus is based upon a misconception
and is without justification. The new genus
Boiotremus to which he attributed 'all tre-
manotids' which have developed tremata during
the whole life of the specimen, i.e. in all onto-
genie stages' is thus a synonym of Tremanotus.

Horny (1963, p. 97) in his discussion of
Tremanotus as redefined by himself, considered
that both the tremata and the flared aperture
were features which developed only at maturity.
However, the development of the tremata was
dependent upon the development of the flared
aperture. This concept of the growth sequence
with its inherent reorganization of the exhalent
system is without justification because of the
presence of tremata throughout the develop-
ment of the entire shell.

Tremanotus pritchardi Cresswell, 1893
(PL 1, fig. 17)

1893 Tremanotus pritchardi Cresswell, p. 42, pi. 8,

fig. 1.
1913 Tremanotus pritchardi Cresswell; Chapman, p.

227.
1916 Tremanotus pritchardi Cresswell; Chapman, p.

79 in part.

Diagnosis: Large form of genus in which the
relationship between major growth rugae and
tremata is variable; numerous fine growth lines
a«nd open tremata are present.
Description: Large planispiral gastropod with
a widely expanded aperture in the final growth
stage; wide umbilici; whorl profile gently arched
dorsally, more strongly curved on the sides
turning sharply into the wide and deep umbilici,
flattened on the inner surface; aperture in final
growth stage sub-oval; neither a sinus nor
tremata are developed on the dorsal surface
of the expanded region of the aperture; pos-
terior to the expanded apertural region, the
existence of a small sinus situated medially on
the dorsal crest of the whorl is indicated by a
slight posterior flexure of the growth lines; along
the resultant selenizone numerous ovoid tre-
mata are developed, the most anterior trema
is represented by a solid protrusion over which
growth lines pass; then follows a number of
open tremata, up to 11, the tremata preceding

C. B. TASSELL

the open trcmata are sealed and flush with the
whorl surface; beween tremata the growth lines
are directed posteriorly towards the earlier
tremata; as well as the closely spaced growth
lines, prominent growth rugae are developed;
the relationship of the growth rugae to the
tremata is variable; in some cases the rugae
intersect the selenizone between trcmata, in
others at the trema; sculpture is composed of
numerous spiral costae which arise in the
umbilici; sculptural elements vary from one to
three orders, sculpture and growth lines form
a reticulate pattern over the entire whorl sur-
face.

Dimensions:

Lap

Wap

Wit

M.U.G.D. 1666
P914

46
43

43
40

35
33

Location of Types: Melbourne University Geo-
logy Department. Holotype, M.U.G.D. 1666
and counterpart M.U.G.D. 1667, G. B. Prit-
chard Coll.

Material: Holotype, counterpart, and 12 other
specimens.

Discussion: This species is distinguished from
the type species, T. alpheus, by having more
open tremata, up to 11 as compared to the
latter's 6 to 8. Transverse growth lines, princi-
pally fine ones of a type lacking in the American
form, are more abundant. The relationship of
the growth rugae and the tremata is quite
variable in the Lilydale form. However, rugae
only occur between tremata in the type species.
The last potential trema is closed, whereas no
mention of this was made by Knight (1941,
p. 354) in his redescription of the type species.
Knight postulated that the tremata were
formed during periods of growth when the
flared aperture was not being deposited. At
such times a shallow sinus in the outer dorsal
lip formed a short slit, subsequently closed
anteriorly by growth of the flared aperture.
The flared aperture was then resorbed, the pro-
minent growth rugae marking the point to
which resorption occurred. This was followed
by the initial growth type in which another
trema was formed. This cycle was repeated
numerous times as is indicated by the number
of tremata.

Material from Lilydale suggests a mode of
growth in T. pritchardi different from that
postulated by Knight.

Initial growth, as in Haliotis, saw the de-
velopment of a short sinus, which with subse-
quent growth of the apertural lips was closed,
so forming a trema. Then followed a period in
which only an exceedingly narrow and shallow
sinus was developed. Subsequently this sinus
deepened and another trema formed. With
growth, earlier tremata were no longer required
and sealed as in Haliotis. This process of growth
continued until the animal reached its penulti-
mate growth stage. At this point the mode of
growth changed. No longer was a sinus formed.
Rather the prominent flared aperture was de-
veloped. Frequently the sculpture present
on the flared aperture is quite different to the
rest of the shell, e.g. all but the first order spiral
sculpture may be absent.

This growth cycle differs from that suggested
by Knight, in that it does not require the flared

PLATE 1

Fig. 1 — Michelia danvini (de Koninck), ANU

36852, hypotvpe, Bloomfield Property, Yass,
N.SAV., XI.

Fig. 2 —Michelia darwini (de Koninck), ANU

36853, hypotvpe, Bloomfield Property, Yass,
N.S.W., XL

Michelia brazieri (Etheridge), P1058, hypo-
type, XI.

— Loxonema australis (Chapman), P38508,
hypotype, XI.

Michelia brazieri (Etheridge), P1059, hypo-
type, XI.

— Belterophon (Bellerophon) cresswelli Eth-
eridge, P12838, hypotype, X 2/3.
•Straparollus {Euomphalus) northi (Eth-
eridge). P28716. hypotype, X 21.

— Straparollus {Euomphalus) northi (Eth-
eridge), P28714, hypotype, X 2/3.

10 — Naticopsis (Naticopsis) lihdalensis Cress-
well, P37740, hypotype, XT.

1-12 — Phanerotrema australis Etheridge, F.
1332, syntype, X H (approx.).

— Scalaetrochus lindstromi Etheridge, F.1137,
holotype, X 2/3.

— Stcnoloron subaequ'tlatera (Chapman),
P37643, hypotype, X 2/3.

: — Scalaetrochus lindstromi Etheridge, F.1137,
holotype X 2/3.

Scalaetrochus lindstromi Etheridge, P38505,
hypotype X 2/3. Basal view showing open
umbilicus.
Fig. 17 — Tremanotus pritchardi Cresswell, M.U.G.D.

1666, holotype, X 2/3.
Fig. 18 — Scalaetrochus lindstromi Etheridge, F.1137,
holotype, X 2/3. Basal view.

Fig.

3 •

Fig.

4 •

Fig.

5 -

Fig.

6 ■

Fig.

7 -

Fig.

8 ■

Figs

. 9-

Figs

. 1

Fig.

13-

Fig.

14-

Fig.

15-

Fig.

16-

MEM. NAT. MUS. VICT. 37 PLATE 1

;,'

REVISION OF THE GASTROPOD FAUNA

aperture to be deposited and resorbed numerous
times. Here the flared aperture is considered
to be a gerontic feature. It is also considered
that the type species mode of growth was the
same as that of T. pritchardL

In his discussion of Boiotremus, Horny
(1963, p. 101) noted the presence of tremata
throughout the entire life of the specimen and
the development of periodically widened aper-
tures. He did not describe the extent and
nature of this 'periodical widened aperture'.
That Tremanotus did not develop a large
flared aperture periodically has been discussed
earlier. Horny did not make mention of resorp-
tion of this widened aperture. The only other
method of removal would be purely mechanical
by abrasion. Whatever the method, the removal
of this region results in the formation of a
feature he termed a 'scar 1 . This is in fact the
same as the major growth rugae observed in
the type species and the species from Lilydale.
As previously noted the distribution and de-
gree of development of these growth rugae
is quite variable amongst members of the same
species and between different species. This is
also true of the species figured by Horny.

Although Horny (1963) figured two speci-
mens NM-L5727 and NM-L5729 (PI. 26,
fig. 2 and PI. 27, figs. 2, 3, 4 and 5) which he
considered show the widened aperture, these
specimens could equally as well be mature
specimens with badly damaged expanded aper-
tural regions. Because of the great variability
in the development of the growth rugae or
'scars' it is considered highly unlikely that the
widened apertures as described by Horny were
developed. To demonstrate their existence un-
equivocally would require a mature specimen
with both the final expanded apcrtural region
and the older widened apertures preserved.
Their appearance would be much the same as
varices of certain gastropods. If, in fact they do
exist, their great variability as reflected by the
growth rugae or 'scars' would make them un-
satisfactory as a generic characteristic.

Chapman (1916, p. 79) mentioned a speci-
men from Marble Creek, Thomson River,
Victoria. Subsequently Talent and Philip (1956)
described a new species T. cyclocostatus from

this locality. This is distinguished from T.
pritchardi by its considerably smaller size and
much finer growth lines and sculpture. It also
has fewer foliaceous growth rugae and those
that are present are irregularly developed.
Subfamily Bellerophontinae McCoy, 1851.

Genus Bellerophon Montford, 1808

Subgenus Bellerophon (Bellerophon) Mont-
fort, 1808

Type Species: Bellerophom vasulites Montfort,
1808; Middle Devonian; The Eifel, Germany.

Bellerophon (Bellerophon) cresswelli Etheridge,
1891

(PI. 1, fig. 6; PI. 3, figs. 3, 4, 5, 6, 9)

1891 Bellerophon cresswelli Etheridgc, p. 130, pi. 19,

figs. 6-8.
1913 Bellerophon cresswelli Etheridge; Chapman, p.

227.
1916 Bellerophon cresswelli Etheridge; Chapman, p.

80, pi. 2, fig. 12, pi. 4, fig. 53.
1916 Bellerophon pisutn Chapman, p. 80 pi. 2, figs.

9-11.

Diagnosis; Typical form of genus with thick-
ened outer lip which extends posteriorly beyond
the umbilical region; broadly crescentic aper-
ture; selenizone only slightly elevated and bor-
dered by two fine threads; sculpture composed
of very line transverse elements.
Description: Medium, subglobular, narrowly
umbilicate planispiral gastropod with broad
involute whorls; whorl profile gently arched
dorsally, more strongly curved on the sides,
turning sharply into the narrow umbilici; aper-
ture very broadly crescentic; margin of outer
lip not flared anteriorly but flared outward in
the lateral and umbilical regions, margin con-
tinues across the parietal wall as a moderately
thick inductura which thickens considerably
towards the lateral margins, greatest thickness
at the junction of the flared outer lip and
parietal wall; narrow, moderately deep slit on
outer lip generating a dorsal selenizone which
is very slightly raised above the whorl surface
and bordered by two very fine threads; sculpture
consists of fine transvere growth lines with
occasional growth rugae, holotype has about
seven whorls.

C. B. TASSELL

Dimensions:

Lap Wap Wh Sw

F.1327
P293

P1087
P12837
P12838
P34938

48
11
41
44
12

37
47
10
44
43
12

3*
37
37

25
41

7-3
40
25*
9#

7 1
6

— 4

— 1

— 4

Location of Types; 1. B. cresswelli, Australian
Museum, Holotype, F.1327. National Museum
of Victoria, Hypotypes, PI 2838, P34938. G.
Sweet Coll.

2. B. pisum, National Museum of Victoria,
Holotype, P1087, A. W. Cresswell Coll.
Material: Holotype, 3 hypotypes and 37 other
specimens.

Discussion: The holotype is partially broken,
revealing the inner whorls which number about
seven. This species is distinguishable from the
similar sized type species B. (B.) vasulites by a
number of features. The aperture of B. (B.)
cresswelli is more broadly crescentic in shape.
The outer lip is considerably thicker in the
Lilydale form and extends further posteriorly
beyond the umbilici. The selenizonc of the
type species is raised further above the whorl
surface and bordered by shallow depressions
rather than the fine threads of the Lilydale
form. Transverse sculpture in B. (B.) cress-
welli is considerably finer, particularly in the
regions near the selenizone.

Chapman (1916, p. 80) referred to a well
preserved specimen in the National Museum
of Victoria collection which exhibits 'a faint
but definite lattice structure of wavy striae
across the growth lines'. However, none of the
specimens examined in this study possess such
ornamentation.

During growth B. (B.) cresswelli changed
slightly in form. The aperture became more
flared. In the juvenile form, flaring was con-
fined to the umbilical region, but with growth
the lateral areas of the outer lip also became
flared. Initially quite thin, the inductura also
thickened with growth, as did the outer lip.
The influence of thickening is quite marked
in the change of shape of the junctions of the
parietal inductura and outer lip. In the juvenile
forms, a marked, moderately deep channel is
present, whereas in the older forms the channel

is considerably shallower. The sculpture also
changes from being quite wide, regular and
nearly foliaceous in smaller forms to generally
closer, finer but more irregular and variable in
the larger forms.

Chapman (1916, p. 80) described B. (B.)
pisum from Lilydale. He did not indicate spe-
cifically how it is distinguishable from this
species. Presumably it is because of its smaller
size, distinct sinus in the outer lip and sculpture
of 'interrupted radial striae . . . between the
lines of growth'. However, none of the features
that Chapman mentioned in his description of
B. pisum are unique. All these features are
found in specimens of B, (B.) cresswelli, par-
ticularly the smaller forms. It is considered that
B. pisum is a juvenile of B. (B.) cresswelli.

Superfamily EUOMPHALACEA de Koninck,
1881

Family Euomphalidae de Koninck, 1881

Genus Straparollus Montfort, 1810

Subgenus Straparollus (Euomphalus) J.

Sowerby, 1814

(= Liomphalus Chapman, 1916)
Type Species: Euomphalus pentangulatus J.
Sowerby, 1814; Lower Carboniferous; near
Dublin, Ireland.

Discussion; Chapman (1916, p. 90) erected the
genus Liomphalus which he distinguished from
Euomphalus in having 'smooth rounded unian-
gulated whorls', and from Straparollus in
possessing a concave spire. The genus is
characterized by the following: discoidal; base
concave; wide umbilicus; spire depressed;
whorls smooth, sometimes with keel; whorls
thicken progressively, in the late stages free
or adpressed.

Knight (1941, p. 174) considered it 'utterly
impossible to arrive at any comprehension of
the genotype species except by comparison
with the adequately described species of other
authors referred by Chapman to his genus'.
Later, Knight (1944, p. 465) placed it in
synonomy with Lytospira Koken.

Philip and Talent (1959, p. 50) demon-
strated that the genus Liomphalus is based
on the internal moulds of Straparollus {Euom-
phalus) northi (Etheridge). Their conclusions

REVISION OF THE GASTROPOD FAUNA

are amply supported by many specimens in the
collections studied.

Straparollus (Euomphalus) northi (Etheridge),

1890

(PI. 1, figs. 7, 8. PL 2, fig. 11. PL 3, figs.

1, 2, 7, 8)

1890 Oriostoma northi Etheridge, p. 64, pi. 9, figs.

6-7.
1894 Oriostoma northi Etheridge, p. 151, pi. 9,

figs. 1-4.
1894 Euomphalus (Oriostoma) northi Etheridge;

Cresswell, p. 157.
1913 Euomphalus northi (Etheridge); Chapman, p.

227.
1916 Euomphalus northi (Etheridge); Chapman,

p. 90.
1916 Liomphalus australis Chapman, p. 90, pi. 4,

figs. 32-33.
1959 Straparolus (Euomphalus) northi (Etheridge);

Philip and Talent, p. 50, pi. 7, figs. 1-12,

pi. 8, figs. 1-2.
1972 Oriostoma northi Etheridge; Yochelson and

Linsley, p. 8, pi. 1, fig. 6, pi. 2, figs. 1-5.

Diagnosis: Dextrally coiled discoidal gastro-
pod; variably developed angulations on upper
and lower whorl surfaces; sculpture initially
consists of strong transverse costae becoming
less developed with growth.
Description: Medium to large discoidal dextral
gastropod; numerous whorls with profile flat to
gently convex and inward sloping between the
upper suture and the variably developed upper
keel at the junction of the upper and outer
whorl surfaces; more than one keel may be
developed on the upper surface; the prominence
of the keel or keels tends to decrease with
increased size; the junction of the outer and
basal whorl surfaces is generally less pro-
nounced than that for the upper surfaces;
sutures impressed; base strongly arched; very
wide umbilicus; spire depressed; aperture
circular; columellar lip thin, concave parietal
lip thin; outer lip slightly thicker and it extends
outwards radially from the upper suture to the
outer whorl surface where a mild concave
flexure is sometimes developed, and then con-
tinues inwards across the base radially to the
suture; no sinus or flexure of growth
lines is developed at the keel, if pre-
sent; sculpture is variably developed on
specimens of different size and on the one
specimen; the smaller specimens and the inner
whorls of the larger specimens possess promi-

nent transverse ridges; on the larger specimens,
fine growth lines gradually succeed the promi-
nent juvenile sculpture; sculpture on the base
while similar to that on the upper and outer
surfaces is less strongly developed; transverse
partitions occur in early whorls; multispiral
operculum of numerous fine whorls of variable
thickness, circular in shape; whorls are normally
visible on the plano-concave exterior surface;
the concave internal surface is smooth being
composed of laminae deposited nearly at right
angles to the opercula rim; the degree to which
the internal surface is concave is quite variable;
a shallow central depression occupies about
one-third of the inner surface; thickness is
variable, even for opercula of the same dia-
meter; operculum fits tightly in the aperture,
being retracted into the shell for about 2 mm.

Dimensions:

Hap

Wap

F.1321a

135

—

F.1139e

20-2

—

P1107

25*

P2503

—

P7609

—

P28373

—

P28498

—

P28499

—

P28707

—

3 +

P28711

—

P28712

—

3 +

P28714

—

P28716

—

P28719

—

P343O0

—

Location of Types: 1. Oriostoma northi, Aus-
tralian Museum. Holotype, F. 1321a, Paratype,
F. 1 139e. National Museum of Victoria. Hypo-
types, P1107, P1115. A. W. Cresswell Coll.
P26890, P28499, P28714, P28716, P28718,
P28719. E. D. Gill Coll. and P34300-34302
which were formerly GSV 55329, 55330 and
55332.

2. Liomphalus australis, National Museum
of Victoria. Holotype, P7609, Paratype, P2503.
A. W. Cresswell Coll.

Material: Holotype, paratype, 13 hypotypes and
49 other specimens.

Discussion: Comparison of S. (£.) northi with
the type species S. (£.) pentangulatus reveals
a number of differences. The former has a
shallower, wider umbilicus and is coiled dex-

C. B. TASSELL

trally. Its upper keel is less strongly developed
and there is an absence of spiral sculptural
elements. The sculpture of the Lilydale form
is more variable; initially the transverse ele-
ments are much stronger than those of the type
species but finally they are considerably weaker.
The sculpture is also less evenly developed on
the upper and basal whorl surfaces.

The dextral coiling of S. (E.) northi is also
a feature of S. (E.) carnicus (Freeh). As
figured by Jhaveri (1969, pi. 21, fig. 8) this
species also possesses strong transverse sculp-
tural elements as does S. (E.) northi. It also
probably possesses an operculum which is very
similar to that of S. (E.) northi, (Yochelson
and Linsley 1972, p. 9).

Yochelson and Linsley (1972, p. 8) de-
scribed the operculum from S. (E.) northi and
compared it with very similar types found in
a number of other genera which belong to more
than one family. Because of this they suggested
implicitly that a revision of these families was
needed. Thus they preferred to leave the species
from Lilydale in the genus Oriostoma as
originally determined by Etheridge. However,
comparison of S. (E.) northi with Oriostoma
barrandei Munier-Chalmas, the type species
reveals sufficient differences to preclude it from
belonging to the latter genus. S. (E.) northi
is discoidal rather than turbiniform and has a
much wider umbilicus. There is an absence of
the spiral sculpture characteristic of the type
species. The form from Lilydale has a more
angular whorl profile and its aperture is more
circular in shape.

Superfamily PLEUROTOMARIACEA Swain-
son, 1840

Family Phanerotrematidae Knight,

1956

Genus Phanerotrema Fischer, 1885

Type Species: Pleurotomaria labrosa Hall,
1860; Lower Devonian; Carlisle, New York,
United States of America.

Phanerotrema australis Etheridge, 1891

(PL l,figs. 11, 12. PI. 2, figs. 3, 10,12)

1891 Phanerotrema australis Etheridge, p. 128, pi.
19, figs. 4-5.

1913 Phanerotrema australis Etheridge; Chapman, p.

227.
1916 Phanerotrema australis Etheridge; Chapman, p.

83, pi. 3, fig. 25.
Diagnosis: Typical form of genus with thick,
short, straight columellar lip, well-developed
parietal inductura and simple rectangular sculp-
ture pattern arising from the intersection of
the collabral growth lines and spiral cords.
Description: Large, turbiniform gastropod with
few whorls; whorl profile sub-angular, gently
arched above and below the selenizone at the
angular periphery; periphery high above mid-
whorl; sutures deeply impressed to sub-canali-
culate; body whorl greatly expanded; umbilicus
absent; columellar lip thickened, continuous
with the thick extensive parietal inductura;
outer lip thin with a broad sinus that forms a
shallow slit at the periphery which gives rise
to the selenizone; from the upper suture to the
selenizone outer lip very gently prosocline;
below the selenizone the outer lip is gently
prosocline; gently concave selenizone mod-
erately wide, and bordered by two threads;
fine collabral growth lines and infrequent
growth rugae, cancellated by two orders of
fine spiral cords to form a rectangular pattern
over the entire whorl surface, occasional speci-
mens have a retrousse intersection.

PLATE 2

-Michelia brazieri (Etheridge), F.1145,

holotype, XI (approx.).
-Gyrodoma etheridgei (Cresswell), F.2542,

hypotype, XI (approx.).
-Phanerotrema australis Etheridge, P.41706,

hypotype, X 2/3.
-Siluriphorus antiquus (Cresswell) , P918,

hypotype, XI. Oblique basal view.
-Scalaetrochus lindstromi Etheridge, P39279,

hypotype, X 11 (approx.). Basal view show-
ing prominent peripheral frill.
-Siluriphorus antiquus (Cresswell), P917,

holotype, XI. Apical view.
-Oriostoma rotundimuratus sp. nov., P1089,

holotype, Apical view.
-Loxonema australis (Chapman), P12851,

holotype, X li.
-Gyrodoma etheridgei (Cresswell), P10187,

holotype, X 1.
-Phanerotrema australis Etheridge, F.39308,

syntype, X 2/3.
-Straparollus (Euomphalus) northi (Eth-
eridge), F.1321a, holotype, X 2/3. Apical

view.
-Phanerotrema australis Etheridge, F.39308,

syntype, X 2/3.

Fig.

2 .

Fig.

5 •

Fig.

7 •

Fig.

8 ■

Fig.

Fig. 12-

MEM. NAT. MUS. VICT. 37 PLATE 2

v.'/"":"-

W 1

'" ' *\

)

'■

:-::,.?:,■:■>

&%§&n >

i»

Sk 1 :

; .

REVISION OF THE GASTROPOD FAUNA

Dimensions:

Ht Wt Hap Wap Wh Clu Cll

F.1332 24 21 — — 4 14 16+

F.39308 82 73 — — 24- — —

P384 63* 47 — — 3+ 29 31

P12841 93 78 — — 4 28* 28*

Location of Types: Australian Museum. Syn-
types, R1332 and F.39308. National Museum
of Victoria. Hypotypes, P 12841, presented by
Dr E. Brooke Nicholls and P41706, A. W.
Cresswell Coll.

Material: Two syntypes, 2 hypotypes and 12
other specimens.

Discussion: P. Australis differs from the type
species in having a straighter, more thickened
and longer columellar lip. The type species'
columellar lip is markedly curved. The parietal
inductura on the form from Lilydale is also
thicker than that of the type species. The
intersection of the collabral growth lines and
spiral sculptural elements in the type species
is retrousse, whereas that of the Lilydale form
is generally considerably simpler.

Family Gosseletinidae Wenz, 1938

Genus Stenoloron Oehlert, 1888
Type Species: Pleurotomaria viennayi Oehlert,
1888; Lower Devonian; Saint-Roch (La Bacon-
nier), department de la Mayenne, France.
Discussion: The presence at Lilydale of a
member of this genus extends the known dis-
tribution as it was previously confined to
Europe and North America.

Stenoloron subaequilatera (Chapman), 1916

(PL 1, fig. 14)

1916 Mourlonia subaequilatera Chapman, p. 83, pi,
3, figs. 18-19.

Diagnosis: Typical form of genus with a seleni-

zone bordered by two cords close to mid-whorl

periphery and finely developed spiral elements

of sculpture.

Description: Medium rotelliform, umbilicate

gastropod; whorl profile well rounded, convex;

moderately impressed sutures; base rounded;

aperture known only in part; outer lip with a

moderately deep and angular sinus that forms

a slit which gives rise to a narrow selenizone;

selenizonc located about one-third of the way

between the mid-whorl periphery and upper

suture; between the upper suture and the seleni-

zone the outer lip is prosocline with a moderate
obliquity; below the selenizone it is prosocyrt,
passing forwards for a short distance before
rounding gently and passing nearly radially
across the base; inner lip not known; seleni-
zone depressed and bordered by two moderately
developed cords; collabral lines strongly de-
veloped; very subdued elements of spiral sculp-
ture.

Dimensions:

P925
P37643

H t Wt Ha p Wap Wh S w

— 5 0-4

— 6 8

22 5

61 — — 6

Location of Types: National Museum of Vic-
toria. Holotype, P925, A. W. Cresswell Coll.
Hypotypc, P37643.

Material: Holotype, hypotype and 2 other
specimens.

Discussion: S. subaequilatera is represented by
only a few specimens, none of which is com-
plete. However, it is possible to distinguish it
from the type species as known on the basis
of Oehlert's original figures and description.
The type species' selenizone is located about
mid-way between the periphery and upper
suture whereas that of the Lilydale form is
considerably closer to the periphery. The form
from Lilydale also possesses two cords border-
ing the selenizone and fine spiral elements of
sculpture. As figured by Oehlert (1888, pi. 9,
figs. 2 and 2a) it is quite possible that the cords
are present although not mentioned in
the description. However, there is no sug-
gestion in Oehlert's figures of the presence of
fine spiral elements of sculpture.

Superfamily ORIOSTOMATACEA Wenz,

1938
Family Ortostomatidae Wenz, 1938

Genus Oriostoma Munier-Chalmas, 1876

Type Species: Oriostoma barrandei Munier-
Chalmas, 1876; Lower Devonian; Bois Roux
quarry at Gahard, near Rennes, France.

Oriostoma rotundimuratus sp. nov.

(PI. 2, fig. 7)
1916 Omphalotrochus zlobosum (Schlotheim); Chap-
man, p. 92, pi. 4, figs. 35-36.

Diagnosis: Small form of genus very similar

C. B. TASSELL

to the type species but with stronger sculpture
and more subdued collabral lines.
Description: Small, low spired, turbiniform gas-
tropod with a few whorls in slight contact;
outer whorl frequently disjunct; whorls increase
rapidly in size, body whorl large; whorl
profile rounded, arching upwards from
the upper suture to the sub-rounded
shoulder, then arching more gently to the
lower slightly less rounded basal angulation
surrounding the umbilicus, finally passing in
a gentle convex arch to the umbilicus; sutures
deep; umbilicate; columellar lip thin; parietal
lip thin; outer lip thick, weakly prosocline;
without sinus or slit; retrousse at the shoulder,
and at each of the elements of spiral sculpture;
sculpture composed of a number of strong
spiral elements of at least two orders; collabral
growth lines fine, slightly foliaceous and
retrousse over each of the spiral sculptural
elements.

Dimensions:

Wt Hap Wap Wh

12 3

7 — — 3

P1089 7

P12850 7

P37644 5

Location of Types: National Museum of Vic-
toria. Holotype, P1089; hypotype, P1088.
A. W. Cresswell Coll.

Material: Holotype, hypotype and 7 other
specimens.

Discussion: Comparison of O. rotundimuratus
with the type species as redescribed by Knight
(1941, p. 219) reveals few differences. The
form from Lilydale has a more rounded whorl
profile particularly in the region of the shoulder.
While the collabral lines of the type species are
more prominent, the spiral elements are
weaker. The type species also possesses a more
arcuate columellar lip and is slightly larger.

Chapman (1916, p. 93) considered this
species to be Trochilites globosus Schlotheim
(1820, p. 162). Comparison of the Lilydale
form with this species is rather difficult, for
Schlotheim's description is brief and he pro-
vided no figures. However, Lindstrom (1884,
p. 162) studied the original specimen from
Gotland upon which Schlotheim based his des-
cription. He synonymized Euomphalus funatus

Sowerby (1823, p. 71) with it.

Comparison of the Lilydale form with illus-
trations of E. funatus indicates considerable
differences between the two. The latter is more
tightly coiled and has a thickened arcuate
columellar lip. It also possesses fewer but
more strongly developed spiral cords. The
Lilydale form has less prominent collabral
growth lines, but these are retrousse over the
spiral elements, a feature apparently lacking
in E. funatus. The sutures of O. rotundimur-
atus are also deeper.

Comparison of 0. rotundimuratus with Om-
phalotrochus globosum (Schlotheim) as re-
described by Lindstrom (1884, p. 160) re-
veals numerous differences. The former is con-
siderably lower spired with deeper sutures
and wider umbilicus. The form from Lilydale
possesses a straighter columellar lip and less
rounded outer whorl profile. It also has less
numerous and weaker spiral sculptural ele-
ments; nor do these elements exhibit a ser-
rated to nodose appearance as in 0. globosum.
The Gotland form also lacks the retrousse
intersection of the spiral and collabral elements
as in O. rotundimuratus.

Superfamily NERITACEA Rafinesque, 1815
Family Neritopsidae Gray, 1847

Genus Naticopsis McCoy, 1844

Subgenus Naticopsis (Naticopsis) McCoy,

1844
Type Species: Naticopsis phillipsi McCoy,
1 844; Lower Carboniferous; Kilcommock,
Longford, Ireland.

Range: Lower Devonian to Triassic. The pre-
sence of a species of this subgenus at Lilydale
extends the lower limit of its range from
the Middle Devonian to the Lower Devonian.

Naticopsis (Naticopsis) lilydalensis

Cresswell, 1893

(PI. 1, fig. 9, 10)
1893 Naticopsis lilydalensis Cresswell, p. 44, pi. 9,

1913 Craspedostoma lilydalensis (Cresswell): Chap-
man, p. 227.

1916 Craspedostoma lilydalensis (Cresswell): Chap-
man, p. 95, pi. 4, fig. 37.

Diagnosis: Form of genus with slight spire,
rounded to only slightly extended base, dis-

REVISION OF THE GASTROPOD FAUNA

tinctly auriform aperture, outer lip basal region
thickened and excavated; slightly arcuate to
straight, thick columellar lip.
Description: Medium, low spired, naticiform
gastropod; whorl profile rounded with upper
surface slightly flattened; shallow adpressed
sutures; base rounded to slightly extended;
without umbilicus; aperture auriform, outer
lip slightly, irregularly, prosocline; moderately
thin on upper and outer surfaces but thickens
considerably on lower surface towards junction
with the columellar lip, thickened region ex-
cavated; columellar lip thick, straight to slight-
ly arcuate; parietal inductura variably devel-
oped, generally moderately thick; collabral
lines fine and closely spaced, occasionally irreg-
ular; infrequently more prominent lines devel-
oped; no other sculpture.

Dimensions:

P948 19

P949 —

P37740 30

Hap

Wap Wh

22-6
29

4
4

Location of Types: National Museum of Vic-
toria. Holotype, P948; Hypotypes P951 and
P37740. A. W. Cresswell Coll.
Material: Holotype, 2 hypotypes and 10 other
specimens.

Discussion: Absence of the apertural region
on the holotype has resulted in some confusion
as to which genus this species should be as-
signed. Chapman (1913, p. 227 ) ascribed
this species to the genus Craspedostoma with-
out reason. Subsequently in 1916, he noted
(p. 95) that the base had an umbilicus and
that 'in a supplementary specimen, a part of
the columellar area of the everted lip is pre-
served, which shows relationship to the above
genus'. The hypotype P951 was considered
by Chapman to possess an umbilicus. This
specimen is in fact an internal mould, the
umbilicus being the space that would have
been occupied by the columella. No specimen
has been found suggesting the presence of an
'everted lip'. Chapman also mentioned the
presence of an obscure cancellated sculpture,
consisting of flattened spiral and collabral ribs.
Again no evidence for the presence of spiral
elements has been found.

Comparison of N. (N>) lilydalensis with the
type species reveals that it is slightly higher
spired but has a less elongate last whorl. The
form from Lilydale also has a more auriform
aperture with a straighter columellar lip. The
lower area of the outer lip is also considerably
thicker. However, the parietal inductura of
the type species is thicker.

Superfamily MURCHISONIACEA Koken,

1896
Family Murchisoniidae Koken, 1896

Genus Murchisonia D'Archiac and De

Verneuil, 1841
Subgenus Murchisonia (Murchisonia)

D'Archiac & De Verneuil, 1841.
Type Species: Muricites turbinatus Schlotheim,
1820; Middle Devonian; Stringocephalus lime-
stone, near Bladbach im Bergischen, Germany.

Murchisonia (Murchisonia) pritchardi

(Etheridge), 1898
(PL 3, fig. 11, 12, 13, 14)

1898 Goniostropha pritchardi Etheridge, p. 71, pi.
15, figs. 1-4.

1913 Murchisonia {Goniostropha) pritchardi Eth-
eridge; Chapman, p. 227.

1916 Goniostropha pritchardi Etheridge; Chapman,
p. 88, pi. 4, fig. 29.

1916 Cyrtostropha lilydalensis Chapman, p. 87, pi. 4,
figs. 26-28.

Diagnosis: Typical form of subgenus but pos-
sessing spiral sculpture above and below the
selenizone.

Description: Medium, high spired, numerous
whorled gastropod with a selenizone between
two prominent cords at the angular periphery;
the whorl face is flat to slightly concave both
above and below the selenizone; sutures mod-
erately deep; base rounded; lacking umbilicus;
columellar lip thin, arcuate and reflexed; junc-
tion of columellar and other lip not known;
parietal inductura thin; outer lip with angular
sinus that forms a slit at the periphery which
generates the selenizone; from the upper suture
to the selenizone the outer lip passes posteriorly
with a moderate obliquity; below the seleni-
zone it passes forwards to the base less
strongly; selenizone concave; collabral lines
fine and weakly developed; sculpture consists
of a number of spiral cords above and below
the selenizone; cords more numerous below the

C B. TASSELL

selenizone than above it; none of the spiral
cords are as strong as those bordering the
selenizone.

Wt Hap Wap Wh Clu C1I

Dimensions:

Ht_

F.4112 20-1 9-4 7 3

F.4112 22-5 9-6 56 4-7 7 3

P935 12-9 7-7 41 3 6 4 3

P936 16-5 9-5 — — 7 3 10

P941 18 7 5 5-6 41+ 6-\ —

P944 31-2 — — — 8+ — —

P946 230 9-5 5-4 40 8 — —

Location of Types: 1. Goniostropha pritchardi,
Australian Museum. Syntypes F.4112. Of
these 5 specimens, one is that figured by
Etheridge, as figure 1, plate 15, another is
that illustrated as figures 2 and 3, plate 15.
The former of these two specimens is the
most complete and is here designated the lecto-
type. National Museum of Victoria. Hypotypes,
P935-940. A. W. Cresswell Coll.

2. Cyrtostropha lilydalensis, National Mus-
eum of Victoria. Holotype, P944. Paratype,
P946. Hypotypes, P941-3, P945, P947.
Material: Lectotype, 4 paralectotypes, 13 hypo-
types and 48 other specimens.
Discussion: Chapman (1916, p.88) distin-
guished C. lilydalensis from G. pritchardi on
the basis of the former's shorter habit, more
angulate whorls and deeper selenizone. How-
ever, these features vary independently of each
other and such variation as does exist is con-
sidered to represent variation within a popu-
lation only.

Chapman (1916, p. 88) suggested that the
specimen of Murchisonia sp. mentioned by
Etheridge (1891, p. 129) might be C. lilydal-
ensis. Re-examination of this specimen indi-
cates that it is not. Rather it is a poorly pre-
served and crushed fragment of what appears
to have been a moderately large specimen of
Michelia brazieri (Etheridge).

M. (M.) pritchardi differs from the type
species noticeably in possessing elements of
spiral sculpture above and below the seleni-
zone. The type species has a slightly more
angular periphery and its growth lines above
the selenizone pass backwards more obliquely.
The Lilydale form has a slightly higher peri-
pheral selenizone.

Genus Michelia Roemer, 1852

Type Species: Michelia cylindrica Roemer,
1854; Devonian; Bockswiese, near Clausthal,
Germany.

Discussion: A new Palaeozoic subgenus of the
Tertiary genus Niso (Risso) was erected by
Etheridge (1890, p. 62). He considered that
ultimately it would 'reveal an organization
differing from Niso in which case I would
propose for it the name Vetotuba'.

Although Etheridge (1890, p. 63) mention-
ed that Niso darwini de Koninck (1876, p.
127) from Yass had an umbilicus similar to
N. (Vetotuba) brazieri, no reference to JV.
darwini was made in the generic description
and discussion. Thus Knight (1941, p. 382)
considered Vetotuba brazieri to be the type
species.

Knight (1944, p. 459) synonymized Veto-
tuba with Coelocaulus (Oehlert). This latter
genus was in turn synonymized by Knight
et al. (I960, p. 1292) with Michelia, as was
Vetotuba itself.

Michelia brazieri (Etheridge), 1890
(PI. 1, fig. 3, 5, PL 2, fig. 1)
1890 Niso (Vetotuba') brazieri Etheridge, p. 62, pi.
8, figs. 4-5, pi. 9, figs. 2-3.

PLATE 3

Fig. 1 — Straparollus (Euomphalus ) northi (Eth-
eridge). P1115, hypotype, X H. Exterior
of operculum.

Fig. 2 — Straparollus {Euomphalus) northi ( Eth-
eridge), P28719, hypotype, X 1£. Basal
view.

Fig. 3. — Bellerophon (Bellerophon) cresswelii Eth-
eridge, P1087, hypotype, X 3.

Figs. 4-6 — Bellerophon (Bellerophon) cresswelii Eth-
eridge, F.1327, holotype, X 1 1/3 (approx).

Fig. 7 — Straparollus (Euomphalus) northi (Eth-
eridge, P28499, hypotype, X 2/3. Section
showing transverse partitions developed in
early whorls.

Fig. 8 — Straparollus (Euomphalus ) northi (Eth-
eridge), F.1139e, paratype, X 1 (approx.).

Fig. 9 — Bellerophon (Bellerophon) cresswelii Eth-
eridge, P34938, hypotype, X 3.

Fig. 10 — Gyrodoma etheridgei (Cresswell), P38504,
hypotype, X 4/5.

Fig. 11 — Murchisonia (Murchisonia) pritchardi (Eth-
eridge), F.4112b, paralectotype, X 2i.

Fig. 12 — Murchisonia (Murchisonia) pritchardi (Eth-
eridge), F.4112a, lectotype, X 2£.

Fig. 13 — Murchisonia (Murchisonia) pritchardi
(Etheridge), F.4112b, paralectotype, X 2£.

Fig. 14 — Murchisonia (Murchisonia) pritchardi (Eth-
eridge), P946, hypotype, X2.

MEM. NAT. MUS. VICT. 37 PLATE 3

\08>

«>

',J»iX

•i

***■'

■

' 6

REVISION OF THE GASTROPOD FAUNA

1894 Niso (Vetotuba) brazieri Etheridge; Cresswell,

p. 158.
1913 Vetotuba brazieri Etheridge; Chapman, p. 227

(in part).
1916 Coelocaulus brazieri (Etheridge); Chapman, p.

86, pi. 3, figs. 20-22.
1916 Coelocaulus apicalis Chapman, p. 87, pi. 3,

figs. 23-24.
1941 Vetotuba brazieri Etheridge; Knight, p. 382,

pi. 46, figs. 3a-c.

Diagnosis: Medium to large, narrowly umbiii-
cate, cyrtoconoid gastropod, with pseudose-
lenizone.

Description: Medium to large, high spired,
cyrtoconoid gastropod; numerous whorls;
whorl profile gently convex to nearly flat;
sutures shallow, impressed; base flatly rounded
with sub-rounded periphery, narrowly umbili-
cate; columellar lip straight otherwise inner
lip unknown; outer lip unknown except that it
gives rise to a pseudoselenizone that is border-
ed by two ridges; sculpture known only in
part; fine collabral lines present on the whorl
base, umbilicus and lower region of the outer
whorl surface; the collabral lines swing back-
wards moderately from the umbilicus and
continue across the base and onto the lower
area of the outer whorl surface, where they
continue backwards a short distance; moder-
ately thin shell; earlier whorls greatly thick-
ened by internal secondary deposits.

Dimensions: All specimens are incomplete.

F.1145

F.1240

—

P1057

234-

—

P1058

P12842

20 -f

—

P12843

19+

—

P37755

Location of Types: 1. Vetotuba brazieri. Aus-
tralian Museum. Holotype, F.1145 (designated
by Knight (1941, p.382)). Paratype, F. 1240a.
National Museum of Victoria. Hypotypes,
P1057, A. W. Cresswell Coll. P12842, J. S.
Green Coll. P12843, G. B. Pritchard Coll.

2. Coelocaulus apicalis, National Museum
of Victoria. Holotype, P1058. Paratype, P1059.
A. W. Cresswell Coll.

Material: Holotype, 1 paratype, 6 hypotypes
and 52 other specimens. Most of the specimens
lack their apical region.

Discussion: Chapman (1916, p. 86) described
the presence of a 'slit band below median line,
feebly concave bounded by threads above and
below'. He considered that PI 2843 exhibited
the slit band. Re-examination of this specimen
suggests that the irregular spiral grooves are
the result of weathering. However, both PI 060
and P37757 quite clearly possess a pseudosel-
enizone. This feature is only found on speci-
mens preserved in the unweathered limestone.
It is not normally found on specimens that
have weathered free.

Chapman (1916, p. 87) erected a new species
C. apicalis and distinguished it from V. braz-
ieri on the basis of its smaller size, smaller
spiral angle, more numerous whorls, particul-
arly in the apical region, and more regularly
cylindrical umbilicus. Variation in these char-
acters in the material from Lilydale is insuffi-
cient to justify the erection of a new species
and is no more than that expected in a single
population.

Comparison was also made between V. braz-
ieri and Niso darwini by Chapman (1916, p.
87). The form from Yass was distinguished
principally by its smaller size and the more
slender apical region. Etheridge also noted that
N. darwini is smaller than the Lilydale form.
Comparison of the two forms is limited by
the nature of preservation of the Lilydale speci-
mens. However, M. brazieri is generally of a
greater size, particularly length, than the Yass
form. M. darwini is also more slender in the
apical region and has a more angular basal
periphery.

M. brazieri as it is presently known differs
principally from the type species in being
cyrtoconoid in shape. Further comparison be-
tween the two is limited because of the rel-
atively poor preservation of both the type
species and the Lilydale species. One specimen
P40619 is cryptomphalous, the umbilical reg-
ion on the base being covered by a swollen
callus deposit. This feature has only been
observed in the one specimen.

Chapman (1907, p. 73) reported M. braz-
ieri from the limestones of Marble Creek,
Thomson River, Victoria. However, this
specimen and material collected more recently

C. B. TASSELL

by Talent & Philip (1956, p. 62) is too poorly
preserved for specific identification.

Michelia darwini (de Koninck), 1876
(PL 1. fig. 1, 2)

1876 Niso darwinii de Koninck, p. 127, pi. 4, figs.

11, lla-c.
1898 Vetotuba darwinii (de Koninck); Dunn, p.

101. English translation of above.
1916 CoeJocaulus darwinii (de Koninck); Chapman,

p. 86.
1941 Niso darwinii de Koninck; Knight, p. 382.

Diagnosis: Small to medium, narrowly umbili-
cate cyrtoconoid gastropod.
Description: Small to medium, high spired,
cyrtoconoid gastropod; numerous whorls; whorl
profile gently convex to nearly flat; sutures
shallow, impressed; base gently rounded with
angular periphery, narrowly umbilicate; aper-
ture subrhomboidal; columellar lip straight
and thin, outer lip thin, straight from the up-
per suture and passing backwards to the basal
edge very gently; sculpture unknown; moder-
ately thin shell.

Dimensions: All material broken.

Ht _Wt Wh Hap Wap

ANU 36851

210

11-7

ANU 36852

31 5

13-2

ANU 36853

35-5

18 6

7-5

PI 2699

9-6

4-4

PI 2700

17 4

5-7

. .

P12701

13 4

7 6

—

Location of Types: 1. Niso darwini, the speci-
mens figured by de Koninck were destroyed
by fire when the Garden Palace in Sydney
was burnt on September 22nd, 1882.

2. Michelia darwini, Geology Department,
Australian National University. Hypotypes,
ANU 36852 and ANU 36853.
Type Locality: 1. Niso darwini, a black com-
pact limestone in the Yass District.

2. Michelia darwini; Chatterton's (1973,
p. 140) locality B, in the lower half of the
'Receptaculites' limestone about 400 m east-
southeast of the homestead on Bloomficld
Property, Parish of Waroo, near Yass.
Stratigraphic Range: The 'Receptaculites' Lime-
stone is considered by Strusz (1972) to be
Emsian.

Material: hypotypes and 23 other specimens.
Discussion: De Koninck (1876, p. 127) gave

the location of his specimens as a black lime-
stone in the Yass District. Etheridge (1890,
p. 63) subsequently gave the location of de
Koninck's specimens as the Upper Silurian,
probably Wenlockian beds near Yass. Speci-
mens from the Shearsby Collection in the
National Museum of Victoria were collected
from Portion 208, Parish of Waroo, N.S.W.
Specimens from the Geology Department, Aus-
tralian National University are from the local-
ities B and M of Chatterton (1973, p.140).
Both these localities are separated from the
Upper Silurian sediments at Yass by a major
geological structure.

The absence of a pseudoselenizone in M.
darwini is another distinguishing feature be-
tween the two species. However, the limited
preservation of this feature in the Lilydale
material suggests its absence at Yass may be
due to the nature of preservation.

Family Plethospiridae Wenz, 1938

Genus Gyrodoma Etheridge, 1898

Type Species: Eunema etheridgei Cresswell,

1893; Lower Devonian; Lilydale Limestone,

Lilydale.

LOCATION OF TYPES
TYPE LOCALITY

Gyrodoma etheridgei (Cresswell), 1893

(PL 2, fig. 2, 9. PL 3, fig. 10)

1893 Eunema etheridgei Cresswell, p. 42, pi. 8, fig. 2.
1898 Gyrodoma etheridgei (Cresswell); Etheridge,

p. 72, pi. 16, fig. 1.
1913 Gxrodoma etheridgei (Cresswell); Chapman,

p. 227.
1941 Gyrodoma etheridgei (Cresswell); Knight, p.

138, pi. 4, figs. la-c.
1960 1 Gxrodoma etheridgei (Cresswell); Knight et

at., p.I296, fig. 192, 2.

Diagnosis: Large, high-spired gastropod with
rounded whorls and deep sutures; selenizone
broad, flat; sculpture, numerous spiral threads;
typically threads also on selenizone.
Description: Large, high spired gastropod;
whorl profile well rounded with deeply im-
pressed sutures; peripheral selenizone at mid-
whorl; minutely umbilicate; inner lip concave;
columellar lip thickened and reflexed extends
as inductura to envelope the umbilicus; pariet-
al inductura thin and extensively developed;
outer lip not known in detail, no growth lines

REVISION OF THE GASTROPOD FAUNA

known; outer lip gives rise to a broad, flat,
sculptured selenizone, bounded by threads
slightly more prominent than those constituting
the remaining whorl sculpture; the selenizone
in the larger whorls generally has a single
fine median spiral thread dividing it into two
equal parts; the selenizone in the earlier whorls
may have up to three threads so dividing it;
sculpture composed of numerous fine spiral
threads of two or more orders above and
below the selenizone; more threads below the
selenizone than above it; threads continue into
the umbilicus.

Dimensions:
Ht

P10187
P38503
P38504
F.2542

Hap

Wap Wh

29+
62
51
62

35 +
31
25
29

— 5+

— 4+

— 5 +

Location of Types: National Museum of Vic-
toria. Holotype, P10187. Presented by the
Rev. A. W. Cresswell to the National Museum
of Victoria on September 9, 1908. Hypotype
P38504. Australian Museum. Hypotype, F.25
42.

Material: Holotype, 2 hypotypes and 24 other
specimens.

Discussion: As figured by Cresswell the holo-
type has vertical growth lines on the divided
selenizone. However, on examination of this
specimen, no suggestion of their presence was
found. Etheridge noted that while the holotype
has a divided selenizone, hypotype F.2542 is
without. Typically the selenizone has one or
more threads developed on it.

Knight (1941, p. 138) designated as the
holotype the left hand illustration in fig. 2, PI.
8 of Cresswell (1893). However, it is con-
sidered both illustrations are of the same speci-
men. This is certainly true for the other
specimens on the plate Tremanotus pritchardi
and Siluriphorus antiquus which are similarly
figured. Comparison of the holotype with the
right-hand illustration suggests that the smaller
whorl of the holotype has been broken sub-
sequently. Supporting this is the presence of
a fresh area of matrix and shell on the upper
surface of the holotype.

Superfamily PSEUDOPHORACEA S. A.

Miller, 1889
Family Pseudophoridae S. A. Miller,

1889
Genus Scalaetrochus Etheridge, 1890

Type Species: Trochus (Scalaetrochus) lin-
strotni Etheridge, 1890; Lower Devonian;
Lilydale Limestone, Lilydale.

Scalaetrochus lindstromi Etheridge, 1890
(PI. 1, fig. 13, 15, 16, 18. PL 2, fig. 5)

1890 Trochus (Scalaetrochus) lindstromi Etheridge,

p. 66, pi. 8, figs. 1-2.
1913 Trochus (Scalaetrochus) Undstroemi Etheridge,

Chapman, p. 228.
1916 Scalaetrochus Undstroemi Etheridge; Chapman,

p. 94.
1941 Scalaetrochus lindstromi Etheridge; Knight, p.

306, pi. 59, figs. 3a-d.

1959 Scalaetrochus lindstromi Etheridge; Philip and
Talent, p. 53, pi. 8, figs. 3-8.

1960 Scalaetrochus Undstroemi Etheridge; Knight
et al., p. 1298, fig. 195, 1.

Diagnosis: Large trochiform low whorled gas-
tropod with mildly concave cryptomphalous
base and narrow peripheral frill; callus deposit
beginning in aperture and filling peripheral
angle; deposit variably developed in umbilicus,
collabral lines moderately prosocline on outer
whorl surfaces.

Description: Large trochiform cryptomphalous
gastropod with mildly concave base; irregular
sutures flush to slightly protruding; whorl
profile gently to moderately concave; periphery
angular, forming narrow frill; columellar lip
thickened and strongly oblique outwards;
parietal inductura thin or wanting; thickened
outer lip moderately prosocline from the upper
suture to the basal periphery, it continues
obliquely across the base to the columellar
Up; the columellar and outer lips on the base
are strongly concave; a callus is variably devel-
oped in the umbilicus, ranging from near
absence in the umbilicus to complete infilling
of the umbilicus; umbilical callus continuous
with the callus which occupies the peripheral
angle; this material is deposited anterior to
the aperture; collabral, growth lines, fine to
slightly foliaceous on the outer whorl surface;
collabral lines on the base are fine and when
the umbilicus is open continue into it; occas-
ional rugae occur on the base.

C. B. TASSELL

Dimensions:

Hap

Wap Wh

F.1137

P294
P921
P16024
P38505

47 72 18 36 5

40 64 — 25+ 6

_ 60 — — —

43 58 — — 6

39 64 — — 5

P38506 36+ 64 15 30 5

Location of Types: Australian Museum. Holo-
type, F.1137. National Museum of Victoria.
Hypotypes, P38505 and P39279. E. D. Gill
Coll.

Material: Holotype, 2 hypotypes and 37 other
specimens.

Discussion: As originally described by Ether-
idge, S. lindstromi was without an umbilicus.
Philip & Talent (1959, p. 53) indicated that
it is cryptomphalous. However, the degree of
development of the callus is more variable
than that intimated by Chapman (1916, p.
93) or stated by Philip and Talent.

Specimens have frequently been crushed,
the base suffering the greatest distortion and
damage. This occurs generally where the
whorl is thinnest, at the inner edge of the
peripheral angle thickening.

Genus Siluriphorus Cossmann, 1918
Type Species: Trochus gotlandicus Lindstrom,
1884; Middle Silurian; the canal near Wes-
toos in Hall, Gotland, Sweden.
Range: Middle Silurian to Lower Devonian.
The presence of a species of this genus at
Lilydale extends the upper limit of its range
from the Middle Silurian to Lower Devonian.
Distribution: Europe and Australia. The pre-
sence at Lilydale of a species of this genus
extends the generic range to include Australia.
Discussion: Philip and Talent (1959, p. 53)
synonymized Siluriphorus with Scalaetrochus
Etheridge. They considered the differences in
size, deflection of the outer lip and sculpture
between the type species are specific rather
than generic.

Close examination of the type species sug-
gests that until there is a review of the entire
family both genera are valid. 5. lindstromi
can be distinguished from S. gotlandicus by
the callus deposit which is not only present
in the umbilicus but also extends to fill the
peripheral angle in advance of the aperture.

The periphery of the Lilydale form is angular
and extends to form a narrow frill, whereas
the Gotland form is characterised by a more
variable periphery. The holotype of Siluripho-
rus gotlandicus has a blunt angular periphery
but in some specimens a blunt frill-like border
is developed. Similarly the collabral lines on
the upper whorl surface of S. gotlandicus are
more variable, the holotype having quite
coarse, strong, irregular imbricating lamellae.
S. lindstromi has very much finer more regular,
foliaceous collabral lines. These lines are also
more arched and are not directed backwards
as obliquely as in the Gotland form. 5. gotland-
icus is characterized by an almost flat to gently
convex whorl profile between obscure shallow
sutures, whereas S. lindstromi has more prom-
inent irregular, flush to slightly protruding
sutures and a distinctly concave whorl profile.

Siluriphorus antiquus (Cresswell), 1893
(PI. 2, fig. 4, 6)

1893 Stomatia antiqua Cresswell, p. 43, pi. 8, fig. 3.

1894 gen. indet. Cresswell, p. 157.

1913 Trochus {Scalaetrochus) antiquus (Cress-
well); Chapman, p. 228.

1916 Scalaetrochus antiquus (Cresswell); Chapman,
p. 93.

Diagnosis: Typical form of genus with sub-
rounded periphery and fine, regular, imbri-
cating growth lamellae.

Description: Medium, trochiform gastropod
with few low whorls and weakly impressed
sutures; whorl profile gently convex to nearly
flat between sutures; basal periphery sub-
angular; base flat to slightly rounded, details
of umbilicus not known but probably cryptom-
phalous; aperture probably avoidal; columellar
lip thickened and strongly oblique outwards;
parietal inductura thin; outer lip strongly ob-
lique backwards from the upper suture to the
periphery, only very weakly prosocline; ob-
liquity maintained across the periphery onto
the whorl base; numerous fine imbricate col-
labral lamellae on outer whorl surface; no
thickening of the peripheral region.
Dimensions:

Hap Wap

P917

14+

39 —

P916

36+

44+ —

P918

13 +

36 —

2+
3+
2+

REVISION OF THE GASTROPOD FAUNA

Location of Types: National Museum of Vic-
toria. Holotype, P917. Hypotype, P918. A.
W. Cresswell Coll

Material: Holotype, hypotype and one other
specimen. All the specimens are incomplete
and crushed to varying degrees.
Discussion: The limited number of specimens
and their crushed state precludes a complete
description of the species.

The type species exhibits considerable vari-
ation in the strength and coarseness of the
growth lamellae, Lindstrom (1884, pi. 14, figs.
1-11). Knight (1941, p. 318) noted that the
holotype is a specimen with coarse, strong
imbricating lamellae. Although cryptomphalous
there is also considerable variation in the form
of the umbilical callus.

S. antiquus in comparison with the holotype
of the type species has weaker, more regular
imbricating growth lamellae on the outer whorl
surface. The type species as exemplified by the
holotype has a more angular periphery than
the Lilydale form. None of the specimens from
Lilydale possesses a bourrelet as found in
some of the specimens from Gotland.

Superfamily LOXONEMATACEA Koken,

1889.

Family Loxonematidae Koken, 1889.

Genus Loxonema Phillips, 1841.

Type Species: Terebra? sinuosa J. deC. Sower-

by, 1839; Middle Silurian; Garden House,

near Aymestry, Shropshire, Britain.

Loxonema australis (Chapman), 1916
(PI. 1, fig. 4, PL 2, fig. 8)

1916 Loxonema sinuosa var. australis Chapman, p.

96, pi. 5, fig. 39.
1949 Loxonema australis (Chapman); Gill, p. 111.
Diagnosis: Medium, intermediate form be-
tween Loxonema and Palaeozygopleura. As
a member of the former genus it possesses a
shallow sinus which deepens slightly with age
and sutures of moderate depth.
Description: Medium, high spired numerous
whorled gastropod with a shallow rounded
sinus in the outer lip; whorl profile gently
arched, periphery at mid-whorl; sutures mod-
erately impressed; base rounded; lacking um-
bilicus; inner lip concave; columellar lip short,

thickened and reflexed; parietal inductura thin;
outer lip with a very shallow wide sinus devel-
oped slightly above mid-whorl; from the upper
suture the growth lines are very steeply pro-
socline to the middle of the sinus, before pass-
ing more gently forwards to the edge of the
base where they curve prosocyrtly to the
centre; sinus deepens slightly with growth; out-
er lip of moderate thickness, except towards
the junction with the columellar lip where
thicker; collabral sculpture consisting of costae
and costellae; protoconch unknown.
Dimensions:

Hap

Wap

P12851

P38507

47+

P3850S

31 +

5+
5+

Location of Types: National Museum of Vic-
toria. Holotype, P12851, J. S. Green Coll.
Hypotype, P38508.

Material: Holotype, hypotype and 8 other
specimens.

Discussion: Comparison of L. australis with
the type species L. sinuosa reveals that the form
from Lilydale has a distinctly shallower and
wider labral sinus. It also has a coarser col-
labral sculpture which weakens slightly on the
base and is slightly larger. There is no evidence
for 'a tendency to form a faint nodose shelf
near the basal part of the whorl', the only
distinguishing feature between the two species
noted by Chapman.

The type species of Palaeozygopleura, P.
alinae Perner, when compared with the Lily-
dale form has a noticeably shallower and
wider labral sinus. It also is considerably
smaller and has much shallower sutures than
L. australis. Thus the form from Lilydale oc-
cupies an intermediate position between these
two genera as regards the depth of the labral
sinus. Such a situation is not unexpected.
Knight (1930) , postulated that Loxonema
gave rise to the Carboniferous Pseudozy-
gopleuridae through an unknown Pseu-
dozygopleura-Yike form that existed in the
Devonian or Lower Carboniferous. This fonn
was most probably represented by the Pala-
eozygopleuridae and more particularly
Palaeozygopleura. Thein and Nitecki (1974,

C. B. TASSELL

p. 29) considered 'that the Palaeozygopleuri-
dae evolved from Loxonema directly during
the Devonian or Mississippian and gave rise to
the Pseudozygoplcuridae that flourished in the
Pennsylvanian and Permian'.

The Mississippian form L. knighti Yochelson
(1962, pi. 17, fig. 11) possesses in its inter-
mediate growth a shallow wide labral sinus
and moderately strong collabral sculpture com-
parable to the intermediate growth stage in
L. auslralis. In its mature form the American
species possesses the characteristically deep
labral sinus of the genus. However, the labral
sinus of the mature Lilydale form does not
deepen as much, although it is deeper than
that of the immature American form. The
species are comparable in size.

Two other genera within the family Loxon-
ematidae also possess shallow sutures, they
being Stylonema and Aulacostrepsis. Both
occur in the Lower Devonian. However, Sty-
lonema is characterized by a very slender shell
and the similar Aulacostrepsis also has a very
small umbilicus.

Although the Lilydale form is obviously in-
termediate between the two genera Loxonema
and Paiaeozygopleura it is for the present as-
signed to the former genus because of its
moderately deep labral sinus, which deepens
slightly with growth, size, moderate sutures
and the nature of the labral sinus in other
genera of the family Loxonematidae.

Gill (1949, p. Ill) in redescribing the holo-
type considered that Chapman s variety was
in fact a new species distinguished by the
coarser nature of the sculpture and the
straighter costellae. He also assigned the form
from Sandy's Creek, Parish of Nungatta, Gipps-
land to this species noting that it is smaller
and that the ornamentation is proportionally
finer. However, Talent (1963, p. 102) con-
sidered that the form from Sandy's Creek
differs sufficiently from L. australis to be con-
sidered a new species, this species being dis-
tinguished by its overall size. Never exceeding
10 mm in width, it is much smaller than L.
australis. The whorls in the form from Sandy's
Creek are also proportionally higher than
those of L. australis. This unnamed species

of Loxonema is also found in the mudstones
at Loyola.

References

Baths, D. E., 1972. A new Devonian crinoid from

Australia. Palaeontology 15 (2): 326-335.
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Victoria, Part 1 Rec. geol. Surv. Vict. 2 (I);

67-80.
, 1913. On the Palaeontology of the Silurian

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1916. New or little-known Victorian fossils

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, 1893. Notes on the Lilydale Limestone.

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-, 1894. Additional notes on the Lilydale

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, 1891. Further descriptions of the Upper

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- 1894. An operculum from the Lilydale

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- 1898. New or little-known Lower Palaeo-

zoic Gastropoda in the collection of the Aus-
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Gill, E. D., 1949. Devonian fossils from Sandy's

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, 1963. Lower Paleozoic Bellerophontina

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-, 1944. In Shimer, H. W. and Shrock, R. R.,

Index fossils of North America. New York, John
Wiley & Sons.

REVISION OF THE GASTROPOD FAUNA

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1960. Descriptions of Palaeozoic Gastropoda. In
Moore, R. C, ed. Treatise on invertebrate paleon-
tology: I. Mollusca (1), Univer. Kansas Press.

Koninck, L. G. de, 1876. Recherches sur les Fossiles
Paleozoiques de la Nouvelle-Galles du Sud (Aus-
tralie). Mem. Soc. Roy. Set. Liege 2, 6. (Trans-
lated, 1898, as: Descriptions of the Palaeozoic
fossils of New South Wales (Australia)). Mem.
geol. Surv. N.S.W., Palaeont. 6.

Lindstrom, G., 1884. On the Silurian Gastropoda
and Pteropoda of Gotland. K. sevenska Vetensk-
Akad. Hand I. 19 (6).

Linsley, R. M., 1968. Gastropods of the Middle
Devonian Anderdon Limestone. Bull. Am.
Paleont. 54 (244).

Manten, A. A., 1971. Silurian Reefs of Gotland.
(Developments in Sedimentology, 13.) Elsevier.

Oehlert, D. P., 1888. Descriptions de quelques
especes devoniennes du department de la May-
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Philip, G. M., 1974. Biostratigraphic procedures and
correlation in the Tasman geosynclinal zone. In
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Schlotheim, E. F. von., 1820. Die Petrefactenkunde
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reibung seiner Sammlung versteinerter und

fossiler Uberreste des Thier-und Pflanzenreichs

der Vorwelt erldutert. Gotha.
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and Wentworth Rivers. Mem. geol. Surv. Vict.

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, and Philip, G. M., 1956. Siluro-Devonian

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Victoria. Proc. R. Soc. Vict. 68: 57-71.
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(Upper Mississippian) Gastropoda of the Illinois

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of Cvclonema Hall (Gastropoda), Bull. Am.

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Yochelson, E. L., 1962. Gastropods from the Red-
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Paleont. 36: 74-80.
, and Dutro, J. T. 1960. Late Paleozoic

Gastropoda from Northern Alaska. Prof. Pap.

U.S. geol. Surv. 334-D.

-, and Linsley, R. M., 1972. Opercula of

two gastropods from the Lilydale Limestone
(Early Devonian) of Victoria, Australia. Mem.
natn. Mus. Vic. 33: 1-13.

A NEW SPECIES OF HEMIERGIS (SCINCIDAE: LYGOSOMINAE) FROM

VICTORIA

By A. J. Coventry

Officer-in-Charge, Herpetology, National Museum of Victoria

Abstract

A new species of lygosomid skink {Hemiergis millewae) is described, from the Victorian
Mallee Region.

Introduction

Field work in the Victorian Mallee since
1973, has revealed the presence of a small,
pentadactyl, lygosomid skink, not previously
known from this State. The species fits into
the genus Hemiergis Wagler, 1830, as recog-
nized by Greer (1967), but does not fit the
only pentadactyl species (//. initiale Werner,
1910). A thorough search of the National
Musuem of Victoria collections brought to light
a further five specimens, which had previously
been identified as Anotis maccoyi (Lucas and
Frost).

The genus Hemiergis was first erected by
Wagler in 1830,and has since been redefined by
many authors, principally Gray (1845), Bou-
lenger (1887), Mittleman (1952), Greer
(1967), Cogger (1975) and Storr (in Press).
Although Cogger and Storr provide the most
recent definitions, Greer's concept of the genus
is accepted here.

Gray provided the first detailed diagnosis of
the genus, restricting it to skinks with trans-
parent lower eyelids, and digits 3-3. He also
considered the presence of paired frontoparie-
tals to be diagnostic, a character which Storr
has shown to be variable within the genus,
even at the specific level. Mittleman restricted
the genus to species having 4-4 or less digits,
thus excluding Werner's 1910 pentadactyl spe-
cies H. initiale. Greer's diagnosis accepted #.
initiale as a true Hemiergis, thus expanding
the genus to include pentadactyl species. Cog-
ger's definition contradicted Greer's in that it
included . . . 'ear opening usually absent, its
position is usually indicated by a slight depres-
sion (a minute opening in one species)'. All
other workers have agreed that the ear open-
ing is in fact covered by scales. Cogger's move

was made to allow the inclusion of Siaphos
maccoyi Lucas and Frost into Hemiergis, thus
making Hemiergis polyphyletic, as Cogger
recognized when he said . . . 'the genus as
recognized here is almost certainly composite,
but relationships are obscure'.

Storr's definition basically agrees with that
of Greer, differing in that Storr said that all
species have a complete series of suboculars,
whereas Greer stated that the subocular row
was 'complete, except in H. initiale'. In addi-
tion to the above characters, Greer also sepa-
rated Hemiergis from Lerista by the presence
of four supraoculars as against 2-4, usually 3,
paired rather than single supradigital scales
on the fourth toe, and non-enlarged as against
greatly enlarged nasal scales.

Genus Hemiergis Wagler, 1830

Hemiergis Wagler, 1830, Nat. syst. amphib.. p. 160.
Type-species Zygnis decresiensis Fitzinger, 1826.

Diagnosis: Small elongate, short-limbed skinks.

Lower eyelid movable, with a transparent disc.

Digits 5-5 to 2-2, normally equal numbers on

fore and hind limbs. Subdigital lamellae less

than 16. Subocular row complete in all non-

pentadactyl species. Ear aperture absent, ear

indicated by a depression. Supradigital scales

paired.

Hemiergis millewae sp. nov.
Fig. 1

Holotype: D47410, adult male in the National Mu-
seum of Victoria, collected at Millewa South bore,
Vic., in 34°56'28"S: 141°4'E on 15.XI.1975 by
A. J. Coventry and P. Mather.

Description: Snout- Vent (S.V.) length 43-5

mm. Length of tail (intact) 75-3 mm, 173%

of S.-V. length. Total length 1188 mm. Length

of hind limb 130 mm, 30% of S.-V. length.

Length of fourth toe 4-7 mm, 36% of hind

A. J. COVENTRY

limb length. No supra or postnasal scales.
Rostral and frontonasal in fairly broad contact.
Frontal and frontonasal in narrow contact. Pre-
frontals large, just failing to meet: contacting
the frontonasal, anterior and posterior loreals,
first supraciliary and frontal. Two loreals,
large and subequal. Frontoparietal entire, in-
terparietal separate, large, almost half the size
of the frontoparietal. Parietals large, barely
contact along the midline. One pair of enlarged
nuchals, followed by a second, single nuchal
on the left hand side. Three enlarged temporals,
the upper largest. Four supraoculars, the
second the largest. Seven supraciliaries, seven
upper ciliaries, the third to fifth largest. Nine
lower ciliaries. Lower eyelid movable with an
extremely large transparent palpebral disc bor-
dered above by the lower ciliaries but otherwise
surrounded by small granular scales.

Length of eye 1-6 mm, length of disc 11
mm, 69% of eye length. Seven upper labials,
the fifth subocular and completely interrupting
the subocular series, of which two are anterior
to, and three posterior to the fifth upper labial.
Seven lower ciliaries. Ear opening completely
covered by scales, indicated by a depression. A
pair of enlarged preanal scales. Limbs short,
pendactyl, when adpressed, failing to meet by
approximately 25% of the distance between
the axilla and the groin. Subdigital lamellae
dark coloured, undivided and smooth, 12 under
the fourth toe. Midbody scales smooth, in 22
rows.

Colour in Life; Uniform dark olive brown dor-
sally, with no trace of spots, striations or lines.
A burnt orange dorso-lateral stripe, approxi-
mately two scales wide, commencing above and
behind the ear, and extending to the hind
limbs. Lateral surfaces off-white, and ventral
surfaces pale yellow. Dorsal and lateral sur-
faces of tail similar to mid-dorsal colour, ventral
surfaces of tail off-white, with darker spots.
Chin whitish, each scale bordered by dark
brown.

Colour in Alcohol: Drab brown dorsally,
whitish laterally, and light grey ventrally. There
is almost no trace of the dorso-lateral stripe.
Paratypes: Fifteen specimens in the National
Museum of Victoria as follows:

1cm

Fig. 1 — Head shields of holotype of Hemiergis mille-
wae D47410, dorsal and lateral views.

D33348 sex undetermined, D33349 male, 0-3 km S
of Millewa South Bore, collected G. Barnes
31.8.1973; the remaining thirteen specimens, all
from Millewa South Bore as follows:

D33341 — 2 sex undetermined, collected by G.
Barnes 29.8.1973;

D38847 sex undetermined, collected A. J. Coventry
4.4.1974.

D40169 sex undetermined, collected A. J. Coventry
24.9.1974;

D47394— 6 males, D47397 female, collected A. J.
Coventry and P. Mather 13.11.1975;

D47398 — 400 females, collected A. J. Coventry and
P. Mather 14.11.1975;

D47418 male, and D47419 female, collected A. J.
Coventry and P. Mather 16.11.1975.

Description of Paratypes; As for holotype ex-
cepting as follows: S.-V. lengths 41*7-58-6
mm (mean 502); hind limb length 11 1-13-7
mm (mean 12-3); percentage of S.-V. length
231-281 (mean 24-7); length of 4th toe
4-5-5-4 mm (mean 4-8) percentage of hind
limb length 35-7-420 (mean 390). Upper

NEW SPECIES OF HEMIERGIS

ciliaries 8-10 (mean 91), lamellae under 4th
toe 12-14 mean (12-8).

Nuchals: one pair (4 specimens), two pairs
(2 specimens), two pairs plus a third on right
side (1 specimen), two pairs plus a third on
left side (1 specimen). D33348 has a damaged
first toe on the left hind foot; D40169 has the
right fore foot missing; D47395 has the second
and third fingers of the right fore limb damaged;
D47397 has a damaged left fore foot, the
second and third fingers, which arise from a
common base, being truncated to appear as a
single digit; D47398 has the third toe on the
right hind foot truncated; D47418 has a dam-
aged fourth toe on the left hind limb.

Colour of all these specimens, in alcohol,
similar to the holotype, excepting that in many
of them, the dorso-lateral stripe is completely
missing.

OTHER SPECIMENS EXAMINED
D1552-3, D1556 from Purnong, S.A.,
Dl 1767-8 from Nonning, S.A., and D47409,
a juvenile, same data as for holotype.
Ecology; Little is known of the ecology of this
species. It resides in porcupine grass {Triodia
sp.) in sandy soil supporting a fairly heavy
cover of mallee scrub. It never appears to
emerge from the Triodia, and was only located
either by burning or the ripping out of this
grass. No activity was observed at any time

of the day or night, although other species of
reptiles (e.g. Ctenotus brachyonyx, Menetia
greyi, Amphibolous barbatus, Amphibolous
fordi, Delma inornata and Lialis burtoni)
were active during the time spent collecting the
type series. Obviously //. millewae is a thig-
motherm, dependent upon Triodia, which one
assumes supplies its food source in the many
forms of invertebrates co-habiting with it.

Key to Hemiergis millewae and other southern
short limbed skinks

1. Ear opening visible, not covered

by scales 2

Ear covered by scales, indicated by

a depression 3

2. Nasals enlarged, meet or almost meet be-
hind rostral Lerista species

Nasals not enlarged Anotis maccoyi

3. Digits 5-5 4

Digits 4-4 or less .... all other known

Hemiergis

4. Hind limb at least 20% of S.-V. length,

subcaudals 12 or more H. millewae

Hind limb less than 20% of S.-V. length,
subcaudals 11 or less H. initiale

This species is named in recognition of the
locality where the type series was collected.

TABLE 1

Showing comparative lengths of hind limbs, and lamellae under longest toe of

H. millewae and its allies

SPECIES

Hind Limb % S.-V. length

Lamellae under longest
toe

Max.

Min.

Mean

No.

Mean

H. millewae
H. initiale
H. peroni
H. decresiensis
A. maccoyi

16
21
18
24
21

29-9
18-7
23-6
16-5
19-5

210
13-2
17-7
111
151

25-1
160
20-2
13-8
17-2

12-14
7-10

10-14
7-8
8-9

12-8
8-8

10-7
7-0
8-3

A. J. COVENTRY

Addendum

Through the courtesy of Dr. T. F. Houston
of the South Australian Museum, I have been
able to examine an additional 119 South Aus-
tralian specimens of this species, which are
under his care. These are as follows: —
R112 Purnong; R3044 A — F near Siam
Station woolshed, 19.3.1950 under Triodia;
R3069 A — Z Birthday Well, Cariewerloo
Station 11.3.1950; R3855 & R1076O — 1 24
km N. Poochera 15.6.1956; R3860, R10767
— 73 & R10775 — 6 Kondoolka Turnoff,
Gawler Ranges 17.6.1956; R5377 & R10762
6 Gawler Ranges March 1963; R10733 —
58 Mamblyn 29.4.1969; R11284 21 km N. E.
Blanchetown 16.2.1969 in sandy soil under a
spinifex bush in mallee scrub; R 12490 A —
Z Miccollo Hill, Siam Station 32° 32' S.: 136°
36' E. 20.4.1971 — ex Triodia Bush; R12619
8 km from Bakara 34° 4' S.: 139° 45' E.
January 1970; R13012 A — G 13 km S
Alawoona36° 6' S.: 140° 32' E. 24.1.1972 —
ex Triodia bushes; R13099 A — C near Oul-
nina homestead, Olary Ridge, 32° 34' S.: 139°
53' E. March — April 1972; R13381 Hiltaba
Reservoir 32° 10' S.: 135° 4' E. 26.8.1972;
R13704 1.5 km S. of Mulgathing Rocks
30° 7' S.: 134° 0' E. 23.4.1973 in Granite
outcrop; R14766 10 km E., 7.5 km N. of
Blanchetown 28.8.1975 low dune mallee.

Of these specimens R3069B, R11284 &
R13012B each have one foetus in utero
R3069M, R13012A & D each have two foe-

tuses in utero, and R3069X, R10736 & R107
48 are hatchlings. This confirms that the spec-
ies is viviparous, having one or two young per
litter, the young being born in late summer or
early autumn. In these series the midbody scale
rows vary from 22 — 24.

Acknowledgements

The author wishes to thank Dr G. M. Storr
of the Western Australian Museum for the
loan of comparative material of Hemiergis
initiate, and for permission to quote from his
unpublished manuscript. Mr P. A. Rawlinson,
Latrobe University for help, encouragement
and critical reading of the manuscript. Mr P.
Mather assisted in both field and laboratory,
and Miss L. Leatham assisted in the laboratory.
Miss R. J. Plant for the drawings of head scales.

Bibliography

Boulenger, G. A., 1887. Catalogue of Lizards in the
British Museum. London.

Cogger, H. G., 1975. Reptiles and Amphibians of
Australia. Reed, Sydney.

Gray, J. E., 1845. Catalogue of Lizards in the British
Museum, London.

Greer, A. E. 1967. A new Generic Arrangement for
Some Australian Scincid Lizards. Breviora 267

Loveridge, A., 1934. Australian Lizards in the Mu-
seum of Comparative Zoology, Cambridge, Mas-
sachusetts. Bull. Mus. comp. Zool. Harw 11' 6

Mittleman, M. B., 1952. A Generic Synopsis of the
Lizards of the Subfamily Lygosominae. Smith-
son, misc. Colin. 117: 17.

Storr, G. M. In press. The Genus Hemiergis Lacer-
tilia, Scincidae in Western Australia.

Werner, F., 1910. Fauna Siidwest-Aust. 2: 480

THE ENDEMIC AUSTRALIAN LIZARD GENUS MORETHIA
(SCINCIDAE; LYGOSOMINAE) IN SOUTHERN AUSTRALIA

By P. A. Rawlinson

Zoology Department, La Trobe University, Bundoora, Victoria, 3083

Abstract

The taxonomy and status of the five southern Australian species of Morethia are discussed
and lectotypes are nominated to stabilize the nomenclature. Details of the species distributions
are provided and the ecology, reproduction, generic relationships and phylogeny of the species
are briefly mentioned.

Introduction

Boulcnger (1887) carried out the first
major revision of the family Scincidae in the
third volume of his Catalogue of Lizards in
the British Museum (Natural History). In the
Preface to this volume Dr A. Gunther stated:
'I feel confident that it will give a fresh
impluse to the systematic study of lizards,
and serve as the standard work for many
years to come 1 . Gunther's confidence was gen-
erally well placed and Boulenger's Catalogues
became the standard reference works. How-
ever, in some groups rather than give a fresh
impulse to systematic studies, Boulenger's
concepts caused a stagnation that lasted more
than half a century. One such group was his
large and clumsy genus Lygosoma with it's
eleven subgenera, and another was the genus
Ablepharus into which Boulenger placed all
skinks with an immoveable transparent lower
eyelid (the 'ablepharine' eye). These genera
were long recognized to be polyphyletic, but
it wasn't until Mittleman (1952) erected the
subfamily Lygosominae and revised the in-
cluded genera that a new stimulus was pro-
vided. Mittleman included the species Boul-
enger had placed in Ablepharus in the Lygos-
ominae and broke up the genus. Since that
time work on the higher taxa of Lygosomine
skinks has made rapid progress and the
species Boulenger included in the genus Ab-
lepharus have been reclassified.

The Australian skinks which were included
in Boulenger's definition of the genus Able-
pharus have been dealt with in two main
papers. Greer (1967) convincingly demon-
strated the artificial nature of the 'ablepharine'
eye taxonomically and phylogenetically when

he united a group of closely related 'able-
pharine' and 'non-ablepharine' skink species
in the genus Lerista. Subsequently Fuhn
(1969) separated the Australian 'ablepharine'
skinks into nine groups, one of which was
the genus Morethia Gray, 1845. Fuhn sep-
arated the genus Morethia on the basis of
skull morphology and in doing so successfully
placed a natural group of closely related
species into one genus. Morethia is now re-
cognized to be an endemic Australian genus
which is not clearly related to any non -Aus-
tralian genus (Storr 1972) and even its
Australian relationships are unclear (Rawlin-
son 1974). The genus presently consists of
six described species, two described subspec-
ies and an undescribed 'race' (Storr 1972).

The Morethia species and subspecies (and
therefore the genus) can be divided into two
geographical and evolutionary groups: a
'northern' group centered in the arid and
semi-arid tropical areas of Australia; and a
'southern' group centered in the arid and
semi-arid temperate areas of Australia. The
'northern' group consists of: M. taeniopleura
taeniopleura (Peters 1874) found in N and
E Queensland, which reaches to 27 i° S but
is centered N of the Tropic of Capricorn
23i° S; M. taeniopleura ruficauda (Lucas
and Frost 1895) found in N Northern Territ-
ory and N Western Australia, which reaches
to 25i° S but is centered N of the Tropic of
Capricorn ; M. taeniopleura exquisita Storr
1972 found in NW Western Australia, which
reaches to 25° S but is centered N of the Tropic
of Capricorn; and an undescribed race of
M. taeniopleura recorded from the N of the
Northern Territory by Storr (1972). These

P. A. RAWLINSON

'northern' taxa are allopatric and appear to
be mutually exclusive (Storr 1972).

The 'southern' group consists of the re-
maining five described species and it appears
to have radiated more widely than the 'north-
ern' group. Three of the five 'southern' spec-
ies, M. adelaidensis, M. butleri and M. ob-
scura, although reasonably widely distributed
are restricted to arid and semi-arid areas S
of 271° S; the fourth species, M. lineoocel-
lata, is virtually restricted to the SW coast
of Western Australia but it extends above the
Tropic of Capricorn to about 20i° S; the
fifth species, M, boulengeri, is widely distii-
buted across S Australia and it extends just
above the Tropic of Capricorn to about 22i°
S in S Queensland. Of the 'southern' taxa,
M. boulengeri and M. butleri are allopatric
and appear to be mutually exclusive as do
M. lineoocellaia and M. obscura; M. butleri
and M. adelaidensis are largely allopatric but
overlap in SE Western Australia; and M.
adelaidensis, M. boulengeri and M. obscura
overlap widely across S Australia. Thus it
can be seen that the Tropic of Capricorn
forms a boundary between the 'northern' and
'southern' groups of Morethia and it is the
five 'southern' species as defined above which
are dealt with in detail in this paper.

Two recent local revisions of the genus
Morethia included the five 'southern' species.
The first revision, published by Smyth (August
31, 1972) was of the South Australian species
but it also included all specimens of Morethia
in the South Australian Museum. Storr pub-
lished the second revision (November 3, 1972),
which was of the Western Australian species
but it also included all specimens of Morethia
in the Western Australian Museum. Unfor-
tunately Symth's and Storr's papers contain
some conflicting interpretations and neither
author examined the types of Morethia lineo-
ocellaia (Dumeril and Bibron 1839) which
was the first of the Morethia species described
and hence the most important taxonomically.
The present paper deals with the 'southern'
Morethia species in Queensland, New South
Wales and Victoria, and as the author has
examined all relevant type specimens, the

opportunity to correct the conflicts between
Smyth's and Storr's papers is taken. Also, as
the author has examined and identified all
Morethia specimens in the Queensland Mus-
eum (QM) Brisbane, the Australian Museum
(AM) Sydney, and the National Museum of
Victoria (NMV) Melbourne, detailed lists of
these specimens are provided under the ap-
propriate headings below to complement the
data in Smyth and Storr. As there is no
Morethia material in the Tasmanian Museum,
Hobart, or the Queen Victoria Museum, Laun-
ceston, the data in Smyth's and Storr's papers
and the present paper represents a complete
listing of the 'southern' Morethia specimens
held in the Australian state museums.

Genus Morethia Gray, 1845

Morethia Gray, J. E., 1845, Catalogue of lizards: 65.

TYPE SPECIES; Morethia anomalus Gray, 1845,
Ibid.: 65 — Ahlepharus lineoocellatus Dumeril,
and Bibron, 1839, Erpetologie Generate 5: 817.

Remarks; Smyth (1972) incorrectly listed
Ablepharus lineoocellatus Dumeril and Bibron
1839 as the type species of the genus. Storr
(1972) listed Morethia anomalus Gray 1845
as the type species by monotypy and in the
same paper designated M . anomalus as a jun-
ior subjective synonym of A. lineoocellatus.
Diagnosis: Small skinks (snout-vent length 17-
56 mm); an 'ablepharine' eye i.e. lower eyelid
an immoveable transparent disc fused to the
eye surface; Frontoparietals and interparietal
fused into a single large shield; parietals con-
tact along midline; supranasal and postnasal
scales present but may be fused to each other
or to nasal scale; frontonasal in broad contact
with the rostral; frontal much larger than the
prefrontals; prefrontals rarely in contact; four
supraoculars, second the largest, first and sec-
ond contact the frontal, second third and fourth
contact the frontoparietal-interparietal shield;
one pair of nuchal scales; seven (occasionally
eight) upper labial scales, the fifth largest
and completely subocular; eight to ten preanal
scales, central four slightly enlarged; limbs
pentadactyl; digits not elongate, 14-27 lamellae
under the fourth toe; body scales smooth,
moderately large, 24-34 rows at midbody;
external ear opening obvious.

THE ENDEMIC AUSTRALIAN LIZARD 29

In the descriptions of Morethia species and — Storr, 1972, /. R. Soc. W. Aust. 55: 73-79.

specimens below, scalation details consistent Flgs " ls 2 '

with the generic description above are not Lectotype: Smyth (1972): ZMB 4733, Zoo-
repeated, logisches Museum der Humboldt Universitat

zu Berlin. Locality: Adelaide. Collector:

KEY TO THE SOUTHERN AUSTRALIAN Schomhun* No other data

SPECIES OF MORETHIA 2>cnomDurgk. JNo otntr data.

1 . Subdigital lamellae acutely keeled, uni- D scnp , lor L u e< f + m / , /' u c ,, .

. °. , . . . a . Remarks: The lectotype, selected by Smyth in

cannate to tricannate; five or srx sup- ln71 +u , '/ i it / i ..

.,- . ' * 9 1971, was the largest of three syntypes under

c i. a- u i i 11 xi ■ " ■•■•'' ' this catalogue number and the paralectotypes

Subdigital lamellae smooth or obtuselv , & u r u 4*kkt*

, . r .... „ have now been given new numbers, ZMB

keeled; six supracihanes 3. .^^ n ~ , , to - , . ,

n t-,. ' ... \ .. A , . , r . 42872-73, data as for lectotype.

2. Five supracilianes, the third, fourth c +u nn7 ^ , c , A^nn\ u fu

* ,J t ' . i j Smyth (1972) and Storr (1972) both coni-

and fifth largest, subequal, and pen- . J A n / , /107 ^ c + u « ~

t t j i t, J th l mented on Peters (1874) use of the name

i j- -x i 1 ii ■ • ' Ablepharus (Morethia) anomalus adelaidensis

subdigital lamellae unicarmate or a -<.u a a n +~ > a • *~~ a~

. . " . . . , , . , and neither considered Peters description ade-

tncarinate M. adelaidensis . TT c ., . , , „ . ,

. .,, . .- ~ ^ ., - quate. However, Smyth considered Peters use

Six .supracilianes, the first the largest Qf ^ mm& constituted a valid indication of

and the remainder forming a deereas- a d ^^ ^ namfi tQ hi aM ^

ing series, junctions of supracilianes cQrdi sekcted Qne of pet£rs , three

with supraocular linear or slightly frQm ^ BerM MmQum as k s

curved, supracilianes do not pene rate ho did no[ ^.^ pete use Qf flie

between supraoculars; subdigital lam- name constituted a vaIid indication of a specie s

ellae unicarmate .... . . . M. feffen ^ he refused tQ ^^ ^ name tQ p eters>

3. First and third supracihanes largest, bu{ he did no{ formall ^ k ^ & mmen
fourth much smaller than third, fourth, mdum Boul (1887) had ascribed the
fifth and sixth successively smaller name ^MaWe,^ to Peters when redescribing

r . .,. , M - boulen 8 en the 'variety' as Ablepharus lineoocellatus C var.

First supracihary never largest, fourth adelaidensis, and there is every chance he ex-

not smaller than third ■;•■.■■■-* amined Peters' types for he stated in the In-

4. Third, fourth and fifth supracihanes traduction to Volume 3 of his Catalogue 'With
largest, subequal and penetrate deeply ^ object of renderine th e account of the
between the supraoculars, sixth much Lacertidae and Scincidae more perfect, I have
smaller than fifth; supranasa often deyoted a mmth tQ the examination of the
fused to nasal . M. Imeoocellala imens in the Berlin Museum'. Storr re-
Fourth supracihary largest, fourth, fifth ded Boule er as the pro er authority for
and sixth form a rapidly decreasing ^ name adel ^ demis and according iy selected
series, third and fourth penetrate deep- a lectot from the spec i m ens described by
ly between supraoculars; supranasal B oulenger in the collection of the British
always separate from nasal. . . M. obscura Museum of Natural History m London: BM

Morethia adelaidensis (Peters 1874) NH 64.10.27.9; Locality: South Australia;

.p. .J -. Collector: G. Krefft; no other data. Smyth had

already examined the specimen which Storr

Ablepharus (Morethia) anomalus adelaidensis Peters, „ n u sprllipnt i u Hp<;ianaterl as lerfntvrvp anH iHpn-

1874, Mher. Preuss. Akad. Wiss.: 375-376. subsequently designated as lectotype and lden-

Ablepharus lineoocellatus C var. adelaidensis Boulen- tified it as M. adelaidensis. The present author

ger, 1887, Catalogue of the Lizards in the British has re _ ex amined Storr's lectotype and confirms

Museum (Natural History). 3: 349. . . , *' t , ,

Ablepharus lineoocellatus (part) Zietz, 1920, Rec. S. it is conspecific with the lectotype selected by

Aust. Mus. 1: 220-221. Smyth (ZMB 4733).

M Mutl6 ad iftits. hT' 1972 ' ^ S ' Smyth's designation of the lectotype is cor-

P. A. RAWLINSON

rect and must be upheld. Storr's action would
have been valid if Peters' use of the name
adelaidensis constituted a nomen nudum, but
Storr did not state this was the case and articles
11., 12. and 16. of the International Code
for Zoological Nomenclature ( 1 964 ) show
Peters' use of the name constituted a valid
indication of the species. It should be noted
that Boulenger himself ascribed the name
adelaidensis to Peters, so both authors were
referring the name to specimens Peters had
examined in the Berlin Museum and there
is a great probability that Boulenger actually
examined Peters' types. Smyth located these
specimens and designated one as lectotype, and
as Smyth's paper was published on August 31,
1972 and Storr's paper was published on Nov-
ember 3, 1972, Smyth's designation predates
Storr's.

Diagnosis: Five supraciliaries, the third, fourth
and fifth the largest, subequal, and all penet-
rate deeply between the supraoculars. Sub-
digital lamellae acutely keeled, unicarinate or
tricarinate. Palmar tubercles elongate and
apically rounded.

Description: Snout-vent length 17-60 mm,
mean 46 1 mm. Total length adults with intact
tails 104-138 mm, mean 118-8 mm. Intact
tail 120-172% of snout-vent length. Suprana-
sals present, widely separated. Postnasals pre-
sent but often fused to supranasals. Prefrontals
narrowly separated. Frontonasal wider than
long. Frontal longer than wide. One to three
ear lobules, usually hidden by projecting prc-
auriculars. Midbody scales in 26-34 rows (us-
ually 28 or 30), mean 29-1. Lamellae under
fourth toe 16-24, mean 19-7.
Colour: Olive-grey to olive-brown dorsal sur-
face, often tinged with red-brown. Small black
spots on back which tend to form broken
lines. Pale dorsolateral stripe occasionally pre-
sent on trunk. Broad dark brown to black
upper lateral stripe strongly speckled with
lighter markings runs from head onto tail.
Wavy edged interrupted white mid-lateral line
runs from upper labials, through ear, above
forelimb and along trunk to hindlimb, usually
margined below by a speckled brown band.
Ventral surface unmarked, white. Males in

breeding condition develop an orange colour
all around the edges of the ventral surfaces
which extends onto the inside surfaces of both
fore and hind limbs and is particularly prom-
inant around the vent and anterior part of the
tail.

Distribution: Arid and semi-arid-areas of SW
Queensland; SW New South Wales; NW Vic-
toria; NE to S South Australia; and SE Western
Australia. (Figure 1.).

Fig. 1. M. adelaidensis

Literature Records: See lists in Smyth (1972)
and Storr (1972).

Specimens Examined: Western A usiralia:
(NMV) Western Australia, R 963; D 1013;
D 1390: 15 mis E of Caiguna, D 44857: South
Australia: (AM) South Australia, 4739:
Fisher, Nullabor Plain, R 7274: (NMV) Cen-
tral Australia, D 1181; D 1183-4: Gawler
Ranges, D 2464-5: Ooldea Well, Overland
Railway, N of Fowlers Bay, D 2473: Overland
Railway, between Ooldea Well and Fowlers
Bay, D2758: Overland Railway to Western
Australia, D 3059; D3109; Lake Eyre, D
3103; D 3128; D 3135: Lake Harry, Birdsville
Track, D 15027: Lake Wangary, Eyre Penin-
sula, D 15054-7; D 15059-60: 3 km NW of
Poonindie, D 15078: Tumby Bay, Eyre Pen-
insula D 15078-80; D 15217: 18i km S of
Maitland, Yorke Peninsula, D 15093-7: Cop-
ley, D 15847-8: Benagerie Station, D 41509,
D 41525: Innamincka area, D 41606-7:
New South Wales: (AM) Moloch, R6445

THE ENDEMIC AUSTRALIAN LIZARD

(3 specimens): Victoria: (NMV) Grampians,
D 1090': 16 km W of Nowingi, D 14652: 13
km S Stawell, D 15064: Kerang, D 15163: 32
km S of Kaniva, D 15174-5: 29 km N of Swan
Hill,D 15177: 10 km W of Nowingi, D 18106.

Morethia boulengeri (Ogilby 1890)

(Hg. 2)

Ablepharus boulengeri Ogilby, 1890, Rec. Aust. Mus.

1: 10-11.

Zietz, 1920, Rec. S. Aust. Mus. 1: 220.

Ablepharus lineoocellatus anomalus (part), Loveridge,

1934, Bull. Mus. Comp. ZooL 77: 377-378.
Morethia boulengeri Smyth, 1972. Rec. S. Aust. Mus.

16: 1-14. Figs. 2, 6.
Storr, 1972, /. R. Soc. W. Aust. 55: 73-79.

Figs. 1, 2.

Lectotype: Smyth (1972): AM R 690, Aus-
tralian Museum, Sydney. Locality: Brawlin,
New South Wales (34° 44' S 148° 02' E).
Collector: H. J. McCooey. No other data.
Description: See Smyth (1972).
Remarks: Smyth (1972) noted that Ogilby
incorrectly recorded a separate interparietal for
the type. Storr (1972) listed Cootamundra,
New South Wales (10 km N of Brawlin) as
the type locality, which is the locality given
for the specimen in the Australian Museum
Register.

Diagnosis: Six supraeiliaries, the first and third
the largest; the fourth much smaller than the
third; and the third, fourth, fifth and sixth
are successively smaller. Subdigital lamellae
smooth or obtusely unicarinate. Palmar tuber-
cles rounded.

Description: Snout-vent length 27-55 mm,
mean 44-6 mm. Total length adults with intact
tails 102-121 mm, mean 109 mm. Intact tail
125-177% of snout-vent length. Supranasals
present, widely separated. Postnasals present
but often fused to supranasals. Prefrontals
separated. Frontonasal wider than long. Front-
al longer than wide. Seven upper labials norm-
ally, rarely eight but the fifth always largest
and entirely subocular. Two to four obtuse
ear lobules (usually two). Midbody scales in
25-32 rows, mean 29-8. Lamellae under the
fourth toe 15-23, mean 19-6.
Colour: Olive-grey to brown dorsal surface,
dorsal scales have 2-5 (usually 3) fine diver-

ging black lines which are often expanded and
merge into black spots or streaks that are
usually distributed irregularly, but may be
organized into interrupted lines or streaks. A
well-defined broad black upper lateral stripe
runs from the eye back to the hindlimb where
it becomes broken up. A prominent moder-
ately wide pure white mid-lateral stripe begins
on the upper labials and runs under the eye
above the forelimb and along the trunk to the
hindlimb. Usually there is also a narrow irreg-
ular black lower lateral stripe below the mid-
lateral stripe. Ventral surface unmarked silver
white. Adult males in breeding condition dev-
elop a bright orange throat. The tail of juv-
eniles is red-orange, not pale fawn as recorded
by Smyth (1972), adults have brown tails.
Distribution: Drier areas of S Northern Terri-
tory; S Queensland; New South Wales; NW
and N Victoria; South Australia; and SE
Western Australia. (Figure 2.).

Fig. 2. M. boulengeri

Literature Records: See lists in Smyth (1972)
and Storr (1972).

Specimens Examined: Northern Territory:
(NMV) Illamurta, James Range, D 473:
Queensland: (AM) Retro Station, Capella, R
12099 (2 specimens); R 12100 (2 specimens);
R 12114 (3 specimens): Mungindi, R 15073;
R 19083: Cunnamulla, R 17121; R 18464:
Gilruth Plains, Cunnamulla, R 20662-3: 8 km

P. A, RAWLINSON

S of Claremont, R 21312: Glenmorgan, R
21317: (NMV) Queensland, D 104: 5 km
W of Amiens, D 9384: (QM) Bell, Darling
Downs, J 2305-9: Retro Station, Capella,
J 6232; J 15768: 24 km W of St. George, J
10456-7: 69 km S of Blackall, J 11564:
Wivenhoe Bridge, Brisbane River Valley,
J 1 1853: 24 km N of Mitchell, J 11854: Bry-
den, J 1 1855: Green swamp, 45 km E of Roma,
J 11856; J 11954: Murphy's Lake, Taroom,
J 11858: 6 km S of Ferndalc, J 11859-60:
Wivenhoe Bridge, 8 km N of Wivenhoe,
J 11862-3: 10 km W of Westgrove HS.,
NW of Gympie, J 11864: li km W of West-
grove HS., NW of Gympie, J 11865: Win
dorah, J 11866-8: Waratah Station via Cun-
namulla, J 11869-71: Alum Rock Station via
Amiens, J 11872: 16 km SE of Capelta,
J 11953: 8 km N of Stanthorpe, J 11955:
Goondiwindi, J 13682: Jandowae, J 13773;
J 13775-6; J 13778-81: Gilruth plains Field
Station, Cunnamulla, J 14353: Bathcaston Sta-
tion, 128 km NW of Marlborough, J 15635-7:
35 km N of Augathella, J 21609: 88 km
NW of Tamboo, J 21610; Binga Station via
Cunnamulla, J 21946: Amiens near Stan-
thorpe, J 22316: Jollys Falls via Stanthorpe,
J 22476: South Australia (AM) Port Lincoln,
R 4764: (NMV) Purnong, D 1216: Overland
Railway to Western Australia, D3058: Why-
alla, D 7743; D 7979-80: Alligator Gorge, Flin-
ders Range, D 15220-2: Wertalona, D 41542:
New South Wales (AM) Cootamundra,
R 687-91; R 950: Brawlin, R 823; R 825-6:
Narrabri, R 1042; R 1058: Western New South
Wales, R 1073-4: Nidgera Ck., near Coot-
amundra, R 1745: Dubbo, R 1750-61: Nar-
romine, R 1780: Boggabri, R 2023; R 2025;
R 4169: near Liverpool, R 2174-5: Gunta-
wang, R 3976: Nevertine to Warren, R 4871:
Buddah Lakes, Trangii, R 5447: Penrose
Farm, Bathurst, R 6268: Moloch, R 6443:
Darling R., between Bourke and Wilcannia,
R 6457: Belubula, R 8077: Jaffa, Ashford
Downs, R 9413: Yaneo district, R 10646:
Yaneo Agricultural School, R 10648: Walcha,
R 10769: Cuddie Springs, Brewarrina, R
11006: Bullerana, Moree, R 11054: Pilliga, R
1 1591 : Bringagee, R 13432: Mootwingie

Watcrholes, R 14678-9: Warrumbungle Moun-
tains, R 14973; R 18921; R 21076: Quam-
bone, R 15120; R 18722; R 18728; R 18753-
4; R 18783: 21 km W of Byrock, R 15212:
24 km W of Byrock, R 15621: Moulamein,
R 15841: Brewarrina, R 17674; R 18727:
Lake Narran, R 17658; R 17711: Bourke,
R 17886: Tocumwal, R 18143-5: Balranald,
R 18146-8: Murray R., Moama, R 20357:
Nymangec, R 20572; R 20642-3: 72 km W
of Cobar, R 27409-10: Round Hill Fauna
Reserve, between Lake Cargelligo and Mt.
Hope, R 27825; R 27834; R 27839-40; R
27862-4; R 27904; R 29669: Old Harbour
Lagoon, near Eumungie, R 28021-2; R 280
24-6: between Tilpa and Barnato, R 28093:
Coonabarabran, R 28095: Duck Ck., 48 km
W of Warren, R 28160: 67 km S of Gilgunnia,
Eubalong Rd., R 28341; R 28347-8: 32 km
S of Cobar, R 29560: Yalgorgrin State Forest,
11 km ex. Gilgandra, R 29645: Lachlan R., 11
km N of Lake Cargelligo, R 30385-7: 64 km
S of Gilgunnia, Mt. Hope to Eubalong Rd.,
R 30391 : Moonabi Lookout, Moonabi Ranges,
R 31741-2: 37 km W of Armidale on Bun-
darra Rd., R 31769; R 31785: 24 km W of
Gilgandra, R 33087-98; R 33100-6: (NMV)
Finley, D 152: Brawlin, D 749-51: Nyngan,
D 10591: Gin Gin, D 10603: Tor Downs, SW
of Menindee, D 10610-1: Moorna, D 12167:
16 km N of Albury, D 14641-2: Savernake
Station, Savernake, D 15043-8; D 15061-3:
Boat Rock Hill, Savernake, D 15065-6:
8 km S of Corowa, D 15067-77: 6i km SW
of Savernake, D 15081: Mulwala, D 15086-7:
Warrumbungle National Park, D 15091: 6i
km S of Morandah, D 15092: 14i km NE of
Urana, D 15130: 3 km NE of Jerilderie, D
15131: 14i km NE of Jerilderie, D 15132:
3 km N of Tocumwal, D 15133: Tocumwal,
D 15134: 10 km N of Cowra, D 15135:
Boree Creek, D 15136-8; D 15142-5: 21 km
SE of Jerilderie, D 15139-41: Sloane, D 15146:
13 km SE of Jerilderie, D 15147: 10 mis
N of Barham, D 15178-92: 90 km W of
Wentworth, D 15850: 24 km NNE of Went-
worth, D 40180-1: Victoria (AM) Browns
Plains, R 3988: (NMV) Murray River, D
917: Ouyen, D 810-1; D 950: Mildura, D

THE ENDEMIC AUSTRALIAN LIZARD

848; D 15218-9: Victoria, D 945: Elmore,
D 1002: Bright, D 1065; D 1067; D 1069:
Tcmplestowe (in error) D 1237: Dimboola,
D 1724; D 2207: Goulburn Valley, D 1808:
Glenrowan, D 1819; D 2517: near Melbourne
(in error) D 2508: Cowangie, D 2717: Gram-
pians, D 3315-6: Kewell, D 3321-3: Red
Cliffs, D 7966; D 7968: Sea Lake, D 9104-5:
Pink Lakes, Linga, D 10713: Birthday Tank,
Sunset Country, D 11185-6; D 33305; D
44865; D 47407: Wood Wood, D 13948-9:
Taminick Gap, Warby Ranges, D 14553-63;
D 14573-5: Warby Ranges, D 14610: Speci-
men Hill, Byawartha, D 14635: 8 km S of
Kiata, Little Desert, D 14911-5; D 14984: 8
km SW of Kiata Little Desert, D 14953: 8 km
SW of Natimuk, D 15042: Walpeup, D 150
49-50: 13 km N of Underbool, D 15051-2:
10 km SE of Wycheproof, D 15053: 13 km
E of Yarrawonga, D 15082: 6i km S of
Nathalia, D 15083: 13 km S of Yarrawonga,
D 15084-5: Kenmere, D 15088: 16 km SW
of Nathalia, D 15089-90: Tutye, D 15097-103:
Pine Plains, D 15104-5: 13 km E of Patche-
wollock, D 15106: Axedale, D 15108: 4 km
E of Mildura, D 15109: Wemen, D 15110:

4 km NE of Wemen, D 15111: 4 km
NE of Mildura, D 15112: 7 km WSW of
Leitchville, D 15113-5: 35 km NW of Nyah,
D 15116: Hattah, D 15117: Mildura Airport,
D 15118-29: Lindsay Point Station, D 15148:

5 km E of Neds Corner Station, D 15149-53;
D 15157-8: 6i km E of Neds Corner Station,
D 15154-5: 6£ km NW of Neds Corner
Station, D 15156: Neds Corner Station, D
15159: Kerang, D 15160-2; D 15164: 5 km
N of Eaglehawk, D 15165-6: 6i km NE of
Wahgunyah, D 15167-72: 6i km E of Tal-
garno, D 15173: 29 km N of Swan Hill, D
15176: Nyah, D 15193: 3 km E of Hattah,
D 15194-206: Tower Hill, Grampian Ranges,
D 15208: 5 km NW of The Crater, Little
Desert, D 15210-3: 6i km N of The Crater,
Little Desert, D 15209: Manangatang, D
15214-6: Tiger Hill via Tatong, D 17710-7:
between Perry and Birthday Tanks, Sunset
Country, D 18064-6: H km E of Birthday
Tank, Sunset Country, D 18076: 7 km W of
Nowingi, D 18102-4: Tank, 6i km E of

Birthday Tank, Sunset Country, D 18165:
SW bank of Rocket Lake, Sunset Country,
D 1 8 1 68 : between Monkeytail and Perry
Tanks, Sunset Country, D 18173: Stockyards
between Perry and Birthday Tanks, Sunset
Country, D 18175-9: Stockyards, Sunset Tank,
Sunset Country, D 18184-5: 5 km NE of
Sunset Tank, Sunset Country, D 18233: 1 km
E of Sunset Tank, Sunset Country, D 18234:
24 km E of Birthday Tank, Sunset Country,
D 33304: 1 km S of Millewa South Bore,
Sunset Country, D 33347: 14i km NE of
Millewa South Bore, Sunset Country, D
33350-1: Millewa South Bore, Sunset Country,
D 33354: Pound Bend, Wemen via Robinvale
D 33358-60: Kulkyne Freehold, D 33528-30:
6i km SE of Pine Plains, D 33536: 5 km E
of Broughton, Nhill Rd., D 33638-43: H km
N of Freuds Plain, Wyperfeld National Park,
D 38799: Main Gate, Wyperfeld National
Park, D 38800-5: 3 km W of Yanac, D 38818:
Sunset Tank, Sunset Country, D 38853;
D 38860-1: 3 km E of Perry Tank, Sunset
Country, D 38856: 3 km N of Bendigo,
D 44834-6: Mildura, D 40141-2: 11 km
WNW of Annuello, D 47343-4: 11 km S of
Boundary Bend, D 47354-6:

Morethia butleri (Storr 1963)
(Fig. 3)

Ablepharus butleri Storr, 1963, W. Aust. Nat. 9:

46-47.
Morethia butleri Smyth, 1972, Rec. S. Aust. Mus. 16:

1-14. Fig. 3.
Storr, 1972, J. R. Soc. W. Aust. 55: 73-79.

Figs. 1, 2.

Holotype: WAM R 20615, Western Australian
Museum, Perth. Locality: Leonora, Western
Australia, 28° 52' S., 121° 23' E. Collectors:
G. M. Storr and R. E. Moreau.
Description: See Storr (1963).
Diagnosis: Six (rarely seven) supraciliaries,
the first the largest and the remainder forming
a decreasing series; junction of supraciliaries,
with supraoculars linear or slightly curved.
Subdigital lamellae acutely keeled and unicari-
nate. Palmar tubercles apically acute.
Description: (After Smyth (1972) and Storr
(1972)) Snout-vent length 25-56 mm. Intact
tail 134-169% of snout-vent length. Suprana-

P. A. RAWLINSON

sals present, widely separated. Postnasals pre-
sent but often fused to supranasals. Prefrontals
separated. Frontonasal wider than long. Front-
al longer than wide. Two to five ear lobules
(usually two or three). Midbody scales in 26-
31 rows (usually 28 or 30), mean 28-9.
Lamellae under fourth toe 19-27, mean 22-4.
Colour: Dorsal surface dark olive-green to
olive-brown sometimes flecked with black, but
usually unmarked. Broad black upper-lateral
stripe may be distinct and well developed or
virtually absent except anteriorally. The white
mid-lateral stripe is equally variable, it may
be distinct and well developed or only readily
distinguishable anteriorally. Lips dark spotted.
Storr (1972) records that the tail (as with M.
boulengeri) is red in juveniles and brown in
adults.

Distribution: Arid and semi-arid parts of S
Western Australia and possibly SW South
Australia S of 27° 30' S. Not known from
any other part of Australia. (Figure 3.).

Literature Records: See lists in Smyth (1972)
and Storr (1972).

Specimens Examined: South Australia (NMV)
Overland Railway to Western Australia, D
2654: No Data (AM) R 38652.

Morcthia lineoocellata (Dumeril and
Bibron 1839)

(Fig. 4)

Ablepharus lineoocellatus (part) Dumeril and Bibron,

1839 Erpetologie Generate 5: 817. Paris.
Morethia anomalus (part) Gray, 1845, Catalogue of

lizards: 65.
Ablepharus lineoocellatus A var. lineoocellatus Bou-

lenger, 1887, Catalogue of the Lizards in the

British Museum (Natural History) 3: 348-349.
Ablepharus lineoocellatus B var. anomalus (part)

Boulenger, 1887 Ibid. 3: 48-349.
Ablepharus lineoocellatus Zietz, 1920, Rec. S. Aust,

Mus. 1: 220-221.
A blepharus lineoocellatus lineoocellatus Loveridge,

1934, Bull. Mus. Comp. Zool. 11: 111.
Morethia lineoocellata Fuhn, 1969, Z. Zool. Svst.

Evolutionsforsch. 1: 67-76. Fig. 7.
Morethia lineoocellata Storr, 1972, J. R. Soc. W. Aust.

55: 73-79. Figs. 1, 3.

Type Series of Ablepharus lineoocellatus,

Dumeril and Bibron, 1839
Remarks: Five specimens (not four as listed
by Guibe, 1954) in the Museum National
d'Historie Naturelle, Paris: MNHP 3092 (old
number 3101). Locality: Nouvelle Hollande.
No other data. Two distinct species are repre-
sented in the type series which was not exam-
ined by either Smyth or Storr. Three of the
syntypes are referable to M. lineoocellata
sensu Storr. The first of these syntypes (the
largest) fits Stores definition of M. lineoocel-
lata in all respects, this specimen is designated
as lectotype below. The second syntype has
a partially separated supranasal scale which
is characteristic of Stores M. obscura, but not
exclusive of M. lineoocellata, this specimen
otherwise fits Storr's definition of the latter
species. The third syntype is aberrant, it has
five supraciliaries owing to the fusion of the
first and second, and it fits Storr's definition
of M. obscura in one respect, for the third
last supraciliary is the largest and the last three
supraciliaries form a rapidly decreasing series.
However, this specimen lacks supranasal scales
which are characteristic of M. obscura. The
remaining two syntypes are conspecific and
distinct from either M. lineoocellata or M.
obscura; both specimens are readily referable
to Menetia greyi Gray 1845 (see below).

In order to restrict the use of the name
Morethia lineoocellata and preserve Storr's

THE ENDEMIC AUSTRALIAN LIZARD

nomenclature, the first syntype listed above
which fits Store's definition exactly is selected
as lectotype here.

Lectotype: MNHP 3092 (Old No. 3101),
Museum d'Historie Naturelle, Paris.
Locality: Nouvelle Hollande (^Australia) .
No other data.

Description: Snout-vent length 42 mm. Tail
(intact) 51 mm, 121% of snout-vent length.
Length of forelimb, 12 mm; length of hindlimb,
18 mm. Snout-axilla, 10 mm; axilla-groin,
24 mm. Six supraciliaries, third, fourth and
fifth subequal in size and penetrating deeply
between the supraoculars; sixth supraciUary
the smallest. Supranasals fused to nasals. Post-
nasals present. One ear lobule. Seven upper
labials, the fifth largest and entirely subocular;
six lower labials. One pair of nuchals. Eight
preanals, four slightly enlarged. Smooth scales,
26 rows round midbody. Subdigital lamellae
smooth, 19 under the fourth toe. Palmar tuber-
cles apically rounded.

Colour: Dorsal surface olive-brown with two
rows of rather indistinct black and white
ocelli. Dorsolateral row of indistinct black and
white ocelli. Black upper lateral band runs
from nostril through the eye, above the fore-
limb and along trunk to tail. Distinct white
mid-lateral stripe runs from upper lip through
ear and above fore and hind limbs to tail.
Black lower lateral band runs below white
mid-lateral stripe, merges into grey lower
lateral region. Ventral surface pure white un-
marked.

Paralectotypes: Four specimens also under
MNHP 3092 (Old No. 3101), Museum d'His-
torie Naturelle, Paris.
Locality: Nouvelle Hollande. No other data.
Paralectotype (a): Snout-vent length 35 mm;
tail (regrown) 41 mm; length of forelimb 11
mm; length of hindlimb 16 mm; snout-axilla
13 mm; axila-groin 20 mm. Scalation as for
lectotype except: supranasal only partially
separated from nasal; 24 rows of scales round
midbody; 18 lamellae under the fourth toe;
ten preanal scales, four slightly enlarged.
Colour as for lectotype except: each of the
two dorsal rows of black and white ocelli have
fused to give continuous white lines margined

by broken black lines on the trunk ad tail;
and on each side the dorsolateral row of
ocellations have also fused to given a con-
tinuous white dorsolateral stripe margined
above and below on the trunk by black. This
specimen is conspecific with the lectotype.
Paralectotype (b): Snout-vent length 41 mm;
tail (regrown) 19 mm; length of forelimb 11
mm; length of hindlimb 17 mm; snout-axilla
9 mm; axilla-groin 20 mm. Scalation as for
lectotype except: five supraciliaries owing to
the fusion of the first and second, second and
third the largest, only the second and third
penetrate deeply between the supraoculars,
and the third, fourth and fifth form a rapidly
decreasing series (this condition which resem-
bles that in M. obscura is apparently caused
by abnormal reduction of the fourth supracil-
iary); 26 rows of scales round midbody; 20
lamellae under the fourth toe; five lower labials.
Colour as for lectotype. Supranasal scales
fused to nasals. If Storr is correct that sup-
ranasals are invariably present in M. obscura,
this specimen is conspecific with the lectotype.
Paralectotype (c): This specimen is neither
conspecific nor congeneric with the lectotype,
and it differs in the following characters:
snout-vent length 29 mm; tail (broken) 5 mm;
length of forelimb 6 mm; length of hindlimb
9 mm. Four fingers and five toes. Scalation
greatly different: all supraciliaries fused into
a single elongate shield; two supraoculars, the
anterior very large and elongate, the posterior
small; 18 scale rows round midbody; 16 lam-
ellae under the fourth toe. Colour as for
Menetia greyi Gray (see Boulenger 1887).
This specimen is conspecific with the syntypes
of Menetia greyi Gray (BMHN X.1.7. a-e;
RR 1946.8.15.1-14 and 1946.8.16.88-99)
and is referable to that species.
Paralectotype (d) : Description as for para-
lectotype (c) except: snout- vent length 27 mm;
tail (broken) 3 mm; length of forelimb 6 mm;
length of hindlimb 9 mm; 18 scale rows round
midbody; 17 lamellae under the fourth toe.
This specimen, is also conspecific with the
syntypes of Menetia greyi Gray (see above)
and is referable to that species.
Type Series of Morethia anomalus Gray, 1845

P. A. RAWLINSON

Remarks: Gray (1845) described this species
from two specimens in the British Museum
collection (BMNH) XI.6a-b; RR 1946.8.15.
74-75). Starr (1972) designated one (BMNH
XL 6b; RR 1946.8.15.75) as lectotype, thus
making M. anomalus a junior subjective syn-
onym of M. Uneoocellata, but did not comment
on the identity of the other syntype. The author
has re-examined Gray's two syntypes and
determined that they belong to different spec-
ies: the lectotype is conspecific with the lecto-
type of M. Uneoocellata which is designated
and described above; but the paralectotype is
conspecific with Storr's (1972) M. obscura
described at the same time that the lectotype
of M. anomalus was designated.
Lectotype: Storr (1972): BMNH XL 6b; RR
1946.8.15.75, British Museum of Natural His-
tory, London. Locality: West Australia. Col-
lector: Mr. Gilbert. No other data.
Description: Snout-vent length 43 mm. Tail
(intact) 63 mm, 149% of snout-vent length.
Length of forelimb, 11 mm; length of hind-
limb, 17 mm. Snout-axilla 9 mm; axilla-groin,

25 mm. Six supraciliaries, third, fourth and
fifth equal in size and penetrating deeply be-
tween the supraoculars; sixth supraciliary the
smallest. Supranasal separate from nasal. Post-
nasals present and separate from supranasal.
One ear lobule. Eight upper labials, sixth the
largest and entirely subocular; seven lower
labials. One pair of nuchals. Eight preanal
scales, four slightly enlarged. Smooth scales,

26 rows round midbody. Subdigital lamellae
smooth, 20 under the fourth toe. Palmar tuber-
cles apically rounded.

Colour: Very faded due to preservation. Dorsal
surface light brown, ocellations not visible.
Light dorsolateral stripe faintly visible. Black
upper lateral band and white midlateral stripe
faintly visible. Ventral surface unmarked,
white.

This specimen is conspecific with the lecto-
type of M. Uneoocellata (MNHP 3092) des-
cribed above. As Storr (1972) did not exam-
ine the syntypes of M. Uneoocellata when he
designated the lectotype of M. anomalus and
placed it in the synonymy of M. Uneoocellata,
his decision was apparently based only on the

written description of the latter species. The
author has examined Storr's lectotype of M.
anomalus and found it to be conspecific with
the lectotype of M. Uneoocellata so Storr's
action is now verified.

Paralectotype: BMNH XI.6a; RR 1946.8.15.
74, British Museum of Natural History, Lon-
don. Locality: West Australia. Collector: Mr
Gilbert. No other data.

Description: Snout-vent length 50 mm. Tail
(broken) 7 mm. Length of forelimb, 12 mm;
length of hindlimb, 19 mm. Snout-axilla, 15
mm; axilla-groin 30 mm. Six supraciliaries,
fourth the largest and fourth, fifth and sixth
form a rapidly decreasing series; the third and
fourth supraciliaries penetrate deeply between
the supraoculars. Supranasals separate from
nasals. Postnasals present and separate from
supranasals. Three ear lobules. Seven upper
labials, fifth largest and entirely subocular; six
lower labials. One pair of nuchals. Eight pre-
anal scales, four slightly enlarged. Smooth
scales, 26 rows round midbody. Subdigital
lamellae smooth, 18 under the fourth toe. Pal-
mar tubercles apically rounded.
Colour: Very faded due to preservation. Dorsal
surface light olive-grey, ocellations not visible-
Black upper lateral band faintly visible. Ven-
tral surface unmarked, white.

This specimen is not conspecific with the
lectotype of M. Uneoocellata, but it is con-
specific with Storr's species M. obscura (see
below) and fits Storr's description of that
species exactly.

Diagnosis: Six supraciliaries (occasionally five
owing to fusion of first and second), the third,
fourth and fifth subequal and penetrate deep-
ly between the supraoculars; the sixth supracil-
iary the smallest. Supranasals usually fused to
the nasals. Subdigital lamellae smooth or
obtusely keeled. Palmar tubercles apically
rounded.

Description: (After Smyth (1972) and Storr
(1972)). Snout-vent length 19-49 mm. Intact
tail 111-247% of snout-vent length. Suprana-
sals normally fused to nasals or separated only
by a shallow or incomplete groove. Postnasals
normally present but only separated from nasal
by a faint groove. Frontonasal wider than

THE ENDEMIC AUSTRALIAN LIZARD

long. Frontal longer than than wide. One to
three ear lobules. Midbody scales in 24-31
rows (usually 26 or 28), mean 27-3. Lamellae
under the fourth toe 16-26, mean 19-7.
Colour: Head coppery brown. Dorsal surface
green, olive-grey or olive-brown, usually mark-
ed with white ocelli outlined in black. Ocelli
may be absent or modified into black or white
spots which sometimes fuse into longitudinal
stripes. White dorsolateral stripe present. Irreg-
ular dark brown or black upper lateral band.
White midlateral stripe margined by black
below usually well developed, runs through
ear, over forelimb and along trunk to hindlimb.
Distribution: On the mainland restricted to
two coastal areas in the SW of Western Aus-
tralia: the mid-west coast from Port Cloates
S to Geraldton; and the lower W coast from
just N of Perth S to Cape Leeuwin and a
short distance inland. Also occurs on islands
of the Montebello Group and Houtmans Abrol-
hos, and from Rottnest and Garden Islands.
Distribution inland very limited. Not known
from any other Australian state. (Figure 4.).

Fig. 4. M. lineoocellato

Literature Records: See list in Storr (1972).
Specimens Examined: Western Australia (AM)
Bunbury, R 30343: (QM) Rottnest Island,
J 12241:

Morethia obscura Storr 1972

(Fig. 5.).

Morethia anomalus (part) Gray, 1845, Catalogue of
lizards: 65.

Ablepharus Hneoocetlatus B var. anomalus (part)

Boulenger, 1887, Catalogue oj the Lizards in the

British Museum (Natural History) 3: 348-349.
Ablepharus lineoocellatus (part) Zietz, 1920, Rec.

S. A ust. Mas. 1: 220-221.
Ablepharus lineoocellatus anomalus (part) l.overidge,

1934, Bull. Mus. Comp. Zool. 11: 377-378.
Morethia lineoocellata Smyth, 1972, Rec S. Aust.

Mus. 16: 1-14. Figs. 4, 6.
Morethia obscura Storr, 1972, J. R. Soc. W. Aust.

55: 73-79. Figs. 1, 3.

Holotype: WAM R 16916, Western Australian
Museum, Perth. Locality: 6 miles cast of Kal-
amunda, Western Australia, 31° 58' S, 116°
08' E. Collector: Mr John Dell. Date of col-
lection: November 7, 1962.
Description: Sec Storr (1972).
Remarks: M. obscura, described by Storr in
1972, is very closely related to M. lineoocel-
lata. Neither Smyth nor Storr examined the
syntypes of M. lineoocellata though both used
the name in their reviews. Smyth recorded
and described specimens from South Australia
under this name. However, he expressed reser-
vations as he noted that South Australian
specimens diflcrcd from many Western Aus-
tralian specimens in several respects viz.: the
invariable possession of supranasal scales; hav-
ing the fifth supraciliary smaller than the
fourth; and having only the third and fourth
supraciliaries penetrate between the suprao-
culars. Smyth noted that Gray (1845) des-
cribed M. anomalus from 'W Australia' and
distinguished it from M. lineoocellata because
the former, but not the latter, had supranasal
scales. Smyth did not use the name M. anom-
alus for South Australian specimens as he
doubted that the presence or absence of sup-
ranasal scales on its own was a good indicat-
ion of a species. He suggested that a careful
study of Western Australian material was
needed as both lineoocellata forms occurred
there and stated that this could result in the
recognition of two species. Storr carried out
such a study and separated the new species
M. obscura from M. lineoocellata using the
condition of the supraciliaries described by
Smyth and the presence of supranasal scales
as the major diagnostic characters. All eastern
Australian specimens, including those described
by Smyth as M. lineoocellata, have proved to
be M. obscura.

P. A. RAWLINSON

The nomenclature of the species presently
recognized in the genus Morethia has now
become very confused, especially in the lineo-
ocellata group. Literature records of M. lineo-
ocellata prior to Smyth and Storr could include
any, or all, of the 'southern' species of More-
thia and should be disregarded unless they
can be verified. Particularly confusing is the
history of the species M. anomalus described
by Gray (1845) from Western Australia. As
mentioned above, Gray distinguished the spec-
ies from A. lineoocellatus because it possessed
supranasal scales. Boulenger (1887) after ex-
amining material in the British Museum which
included Gray's types of M. anomalus, applied
the name Ablepharus lineoocellatus anomalus
to specimens with supranasal scales. Authors
from that time mainly followed Boulenger (e.g.
Loveridge, 1934 and used the name anom-
alus for the more eastern populations of M.
lineoocellata which possess supranasal scales
(i.e. Storr's M. obscura), though, as Smyth
notes, most authors were probably also includ-
ing M. adelaidensis and M. boulengeri under
this name. As recorded above, there were two
syntypes of Gray's M. anomalus, the lectotype
(BMNH XI. 6b, RR 1946.8.15.75) which is
conspecific with M. lineoocellata sensu stricto,
and the paralectotype (BMNH XI.6a, RR
1946.8.15.74) which is conspecific with Storr's
new species M. obscura. The name M. anomala
could have been retained if the paralectotype
had been nominated instead as lectotype for
the taxon. This would also have preserved
recent usage of the name. It is unfortunate
that the name anomalus has been placed in
the synonymy of M. lineoocellata and a new
name, M. obscura, has been introduced. It
must be stressed again that in Smyth's review,
the name M. lineoocellata was applied to speci-
mens which now properly belong in Storr's
species M. obscura and all Smyth's descriptions
etc. apply to this latter species.
Diagnosis: Six supraciliaries (rarely five owing
to the fusion of the first and second), fourth
the largest, and the fourth, fifth and sixth form
a rapidly decreasing series, the third and fourth
(and rarely the fifth) penetrate between the
supraoculars, sixth the smallest. Supranasals

invariably present. Subdigital lamellae smooth
or obtusely keeled and unicarinate; palmar
tubercules apically rounded.
Description: Snout-vent length 1 8-56 mm,
mean 43 mm. Total length of adults with intact
tails 107-129 mm, mean 117 mm. Intact tail
120-189% of snout-vent length. Supranasals
and postnasals always present but often fused
to each other or only separated by a shallow
groove. Supranasals widely separated. In most
specimens the third and fourth supraciliaries
penetrate between the supraoculars, the fourth
is largest, and the fourth, fifth and sixth are
successively smaller; but rarely the fifth sup-
raciliary is nearly as large as the fourth and
it also penetrates between the supraoculars.
Frontonasal wider than long. Frontal longer
than wide. One pair of nuchals. One to four
ear lobules. Midbody scales in 24-31 rows
(usually 26 or 28), mean 27-7. Lamellae
under the fourth toe 14-23, mean 190.
Colour: Olive-brown to olive-grey dorsal sur-
face, usually with dorsal ocellations which
consist of a single scale with the middle third
white and the outer thirds black. The dorsal
ocellations are rarely bold or numerous, and
may be reduced to black flecks or be absent
altogether. Occasionally there is a trace of a
pale dorsolateral stripe. Broad irregular black
upper lateral stripe. Narrow pale irregular mid
lateral stripe usually present, running from eye
through ear over forelimb and back to hind-
limb. The upper lateral and mid lateral stripes
are not as even or bold as those in M. bou-
lengeri.

Distribution: Arid and semi-arid areas of SW
New South Wales; NW Victoria; S South
Australia and offshore islands; and S Western
Australia. (Figure 5.).

Literature Records: See lists in Smyth (1972
as M. lineoocellata) and Storr (1972).
Specimens Examined: Western Australia:
(AM) Perth, R 2457; Western Australia, R
6483; Bornham, R 7689; Cranbrook, R 7690;
Merredin, R 9152; Eradu near Geraldton, R
9164; Woodlands, Tambellup, R 11121; R
11666; Northam, R 12350: South Australia:
(NMV) Purnong, D 1546-51, D 3076; Lake
Wangary, Eyre Peninsula, D 15058: New South

THE ENDEMIC AUSTRALIAN LIZARD

Wales: (AM) Nymangee, R 17675; Round
Hill Fauna Reserve between Lake Cargelligo
and Mt. Hope, R 27860; R 27869; R 27823-4;
R 27833; R 27838; R 27883-4; R 29670-1;
8 km W of Nymangee, R 18481: Victoria:

Fig. 5. M. obscura

(NMV) Raak Plains D 698; Ouyen, D 1040;
D 2416; Red Cliffs, D 7969; Kiata, Little
Desert, D 8958; Pink Lakes near Ouyen,
D 9473-4; Kooloonong, D 13951; Broughtons
Waterhole, Little Desert, D 14917-29; li km
NE of Broughtons Waterhole, D 14930-1;
D 14937; 8 km S of Broughtons Waterhole,
D 14932-3; D 14935; D 14942-3; D 14945; D
14947-8; H km E of Broughtons Waterhole, D
14934; D 14936; D 14940; D 14944; D 14946;
H km W of Broughtons Waterhole, D 14938;
5 km E of Broughtons Waterhole, D 14939;
3 km NNE of Broughtons Waterhole, D 14941;
Stan's Camp, Little Desert, 22 i km SW
of Nhill, D 14949-50; 16 km N of Goroke,
Little Desert, D 14951-2; 14i km S of Murray-
ville, D 18235; Spring, 48 km S of Murrayville,
D 18236; SA-Vic. border due W of Telopea
Downs, Big Desert, D 38819; Red Bluff, Big
Desert, D 38822; D 40174-5; The Springs,
Big Desert, D 38831; 15 km E of Broughton,
NW of Nhill, D 44866: 6 km S of Dimboola,
D 44867: Little Billy Bore, Big Desert, D
47392-3: No accurate data (AM) R 39455.

Ecology

Detailed laboratory studies have been made
only on M. boulengeri. Field observations show
M. adelaidensis and M. obscura to be similar
to M. boulengeri and, from the distributions
of M. butleri and M. lineoocellata, it is con-
sidered that they would also be broadly similar.

M. boulengeri is a heliothermic, insectivorous
skink. The species has high thermal prefer-
ences compared to other skinks from other
temperate areas (Rawlinson, 1974 a,b; 1975).
The voluntary minimum temperature is
29-95°C, the mean preferred temperature is
3409°C, and the voluntary maximum temp-
erature is 39-35°C.

Habitat preferences have not received any
detailed attention. However, it is possible to
state that all species live in open vegetation
forms ranging from semi-desert to woodland.
Morethia species are generally restricted to
areas where mean annual rainfall is less than
50 cm.

Reproduction

M. adelaidensis, M. boulengeri and M. obscura
are all oviparous, but details are known only
for M. boulengeri. Unlike the majority of
lygosomid skinks from cool temperate areas,
this species does not show obligatory sperm
storage overwinter (Rawlinson, 1974 b).
Ovarian and testicular activity commences in
early spring (September to October) and
ovulation occurs in late October to early Nov-
ember. Copulation and fertilization also occurs
at this time. The fertilized eggs are retained
in the oviducts until late January or early
February when they are laid in an advanced
state of development. Clutch size varies from
3 to 5 with a mean of 3-5 (11 observations).

Relationships

There is no doubt that the five southern
species of Morethia are all closely allied and
that with the northern (M. taeniopleura) group
they form a good genus. The relationships of
the genus have been discussed several times
recently (see Greer 1974 and included authors).
Greer recorded that the Morethia species have
an 'alpha' lygosomid palatal bone pattern that

P. A. RAWLINSON

fits them into his 'Group II* of Leiolopisma-like
genera. On this basis he concluded that the
closest Australian genera are Anotis, Crypt-
oblepharus, Emoia, Leiolopisma and Pseu-
demoia (though he included the latter genus
in Leiolopisma) and the present author agrees
fully with this finding. However, using morpho-
logical criteria, Greer went on to construct
a phylogeny in which the 'Group IF Leiolo-
pisma-likQ genera were derived from an an-
cestral species close to Pseudemoia spenceri
(which he placed in the genus Leiolopisma).
Greer considered the genus to be the end of
an evolutionary line that ran from a Pseudem-
oia spenceri-like. ancestor through Emoia to
Cryptoblepharus then Morethia, In a second
lineage Greer considered that the genera Anotis
and Proablepharus arose from Leiolopisma via
the same Pseudemoia spenceri-\\ke ancestor.
The present author considers this phylogeny
to be unlikely as it involves the evolution of
five oviparous genera (Anotis, Cryptoblepharus
Emoia, Morethia and Proablepharus) through
two groups (Pseudemoia and Leiolopisma) that
are placental, viviparous forms. Until full
details of the biology and morphology of all
'Group II' Leiolopisma-like species are known,
their origins, relationships and phylogeny must
remain doubtful.

Discussion

The five southern species of Morethia pre-
sent an interesting pattern of speciation in the
temperate arid and semi-arid areas of Australia.
However, until further details of the ecology
of the various species are known, it is not
possible to meaningfully comment on their
biogeographic and evolutionary significance.
When these details are known, the group will
provide a radiation pattern to compare and
contrast with other related lygosomid groups,
the more mesic temperate genera Lampropho-
lis, Leiolopisma and Pseudemoia, and the
tropical genera Anotis, Cryptoblepharus and
Carlia.

Acknowledgements

The author thanks Dr H. G. Cogger of
the Australian Museum, Mr A. J. Coventry

of the National Museum of Victoria, and Ms
J. Covacevich and Mr G. Ingram of the
Queensland Museum, for help in locating
specimens in the collections under their care.
Ms A. Grandison, Dr E. Arnold and Mr A.
Stimson of the British Museum of Natural
History in London, Professor J. Guibe of the
Museum National d'Historie Naturelle in Paris
and Dr G. Peters of the Zoologisches Museum
der Humboldt in Berlin all gave invaluable
assistance in the location and examination of
type specimens and associated data. Finally
the author wishes to express sincere appreci-
ation for the help given by various staff mem-
bers of the National Museum of Victoria, espe-
cially to Mr A. J. Coventry and Mr T. A.
Darragh for assistance with the manuscript
and Mr J. McNally for allowing the facilities
of the Museum to be used during the collect-
ion of data.

References

Boulenger, G. A., 1887, Catalogue of the Lizards in
the British Museum (Natural History) 3: 3 18-
349.

Dumeril, A. M. C. and G. Bibron, 1839. Erpeto-
logie Generale 5: 817. Paris.

Fuhn, J. E., 1969. The 'polyphyletic' origin of the
genus Ahlepharus (Reptilia, Scincidae) : a case of
parallel evolution. Z. zool, syst. Evolutionsforsch
7: 67-76.

Gray, J. E., 1845. Catalogue of the specimens of
lizards in the collection of the British Museum:
65. British Museum, London.

Greer, A. E., 1967. A new generic arrangement for
some Australian scincid lizards. Brevoria No.
267: 1-19.

, 1970. A subfamilial classification of scincid

lizards. Bull. Mus. Comp. Zool. Harv. 139:
151-183.

; — , 1974. The generic relationships of the

scincid lizard genus Leiolopisma and its relatives,
Aust. J. Zool. Suppl. Ser. No. 31: 1-67.

Guibe, J., 1954. Catalogue des types des lezards.
Colas, Bayeux O.P.I.A.C.L., Paris.

Loveridge, A., 1934. Australian reptiles in the Mu-
seum of Comparative Zoology, Cambridge, Mas-
sachusetts. Bull. Mus. Comp. Zool, Harv. 77:
377-378.

Lucas, A. H. S. and C. Frost, 1895. Preliminary
note of certain new species of lizards from Cen-
tral Australia. Proc. R. Soc. Vict. 7: 264-269.

Mittleman, M. B. 1952. A generic synopsis of the
subfamily Lygosominae. Smithson. Misc. Collect
117: 1-35.

Ogilby, J. D., 1890. Redescription of an Ahlepharus
from Australia. Rec. Aust. Mus. 1: 1-10.

Peters, W., 1 874. Uber einige neue Reptilian
(Lacerta, Eremias. Diploglossus, Euprepes, Lygo-

THE ENDEMIC AUSTRALIAN LIZARD

soma, Sepsina, Ablepharus, Simotes, Onychoce-
phalus). Sber. Dt. Akad. Wiss. Phys. — Math.
Klasse. Juni 1874: 367.

Rawlinson, P. A., 1974a. Biogeography and ecology
of the reptiles of Tasmania and the Bass Strait
area. Ch. 11. In Williams, W. D. (Ed.), Bio-
geography and Ecology in Tasmania. Mono-
graphic Biologicae 24: 230-269. The Hague, Junk.

, 1974b. Revision of the endemic south-
eastern Australian lizard genus Pseudemoia
(Scincidae: Lygosominae). Mem. natn. Mus.
Vict. 35: 87-96.

-, 1975. Two new lizard species from the

genus Leiolopisma (Scincidae: Lygosominae) in

southeastern Australia and Tasmania. Mem.

natn. Mus. Vict. 36: 1-15.
Smyth, M., 1972. The genus Morethia (Lacertilia,

Scincidae) in South Australia. Rec, S. Aust. Mus.

16: 1-14.
Storr, G. M., 1963. Ablepharus butleri, a new scincid

lizard from Western Australia. W. Aust. Nat. 9:

46-47.
, 1972, The genus Morethia (Lacertilia, Scin-
cidae) in Western Australia. /. R. Soc. West. Aust.

55: 73-79.
Zietz, F. R., 1920. Catalogue of Australian lizards.

Rec. S. Aust. Mus. 1: 181-228.

A CREEPING CTENOPHORAN (PLATYCTENEA: CTENOPHORA)
FROM VICTORIA, AUSTRALIA

By Brian J. Smith and Rhyllis J. Plant
Invertebrate Department, National Museum of Victoria, Melbourne

Summary

auu f ? P ?o&J ens ° f 1 a P[ at y c , tei ? e an ctenophoran tentatively identified as Coeloptana wtlteyi,
Abbott, 1907, are described living on red and green algae at the southern end of Port Phillip
Bay, Victoria. These constitute the first record of this group of animals from southern Australia
and only the second record for Australia,

Introduction

During survey work at the southern, end of
Port Phillip Bay, Victoria, Mr Phil Hollis of
the Underwater Research Group of Victoria
discovered several specimens of a small creeping
ctenophoran. These were discovered on detailed
examination of minute algal faunas in an
aquarium following collection of bottom
growths from selected deep water areas in the
Bay, and were brought in alive to the Museum
where they were observed for several days.
They were readily identified as platyctenean
ctenophorans belonging to the genus Coelo-
plana by their creeping habit, the presence of
two retractile pinnate tentacles, a central dorsal
statocyst and the absence of swimming comb-
plates.

This constitutes the first record for the Order
Platyctenea for southern Australia and is only
the second record for Australia, the other being
for the Great Barrier Reef (Stephemson, 1931).
Apart from an otherwise unpublished record
by Dayton and Robillard for Antarctica 1968
(pers. comm. in Gordon, 1969) it is the most
southerly record for this group of unusual
animals.

MATERIAL

Four specimens were discovered on algal
growth taken from 15 5 m in a tidal hole
1-5 km north of Portsea in the southern part
of Port Phillip Bay, Victoria (38° 19' 5; 144°
45' E). All specimens were found by Mr Phil
Hollis, the first in December 1972 and three
more in January 1973. The first and second
specimens were found crawling on Caulerpa sp.
while the others were on red algae. The samples
in each case consisted largely of algae though

some ascidians and other sessile animals were
also present in small numbers. It is therefore
not known with certainty on what substratum
the animals were originally taken. However,
they are not confined to one type of substratum
and appear to be able to readily pass from
one to the other.

Three specimens are preserved in 5% neutral
formalin in the National Museum of Victoria,
Reg. No. G2649.

DESCRIPTION

The animal has a very flat flexible body
capable of extension in any direction, with a
thicker central dome-like region. In the centre
of this domed region is a statocyst, composed
of a central granule suspended in a vesicle,
which is overlaid by a section of the body wall.
There is an aperture in the outer body wall,
which is figure-8 shaped with a central constric-
tion, the long-axis of the aperture being per-
pendicular to the inter-tentacular axis. This
aperture can be opened and closed very rapidly,
presumably by a sphincter muscle system.
Surrounding the statocyst in the central area
are several clear vesicles or pustules. On the
few specimens examined these pustules vary in
number from 8 to 25, ranging in size from
three times as large as the vesicle containing
the statocyst to approximately half its size,
and in organization from a regular arrangement
of radiating lines around the statocyst to a
totally irregular arrangement.

There are two long pinnate tentacles capable
of extension to 6 to 8 times the body diameter.
These tentacles are completely retractable into
two tentacular sheaths at opposite sides of the
body in the central domed region. When the

BRIAN J. SMITH and RHYLLIS J. PLANT

tentacles are retracted the positions of the
tentacular sheaths are barely discernible as
slightly raised, smaller areas of the central
dome region.

The overall body colour is a dark pink to
orange-red with small pale or transparent areas
and some white blotches. This appears to be
made up of small red pigment spots and some
white pigment spots in a largely transparent
general body structure. Contraction of the body
causes an intensification of the colour.

The ventral surface is flat, with less colour,
giving the body a semi-transparent appearance.
The general distribution of the main body or-
gans can be seen through the ventral surface.
The position of the tentacular sheaths can be
seen and also a series of canals surrounding a
meridional canal.

BEHAVIOUR

When crawling the animals expanded to 12
mm in diameter and the tentacles were capable
of extending to at least six to eight times the
body diameter. The animals were first ob-
served crawling on the surface of green or
red algae. Tentacles were streamed either to-
gether or independently and retracted intermit-
tently. The animal crawled freely over the
surface of the alga with no particular part
leading. However, several independent obser-
vations were made of the body being bent
round so that the two tentacular sheaths were
positioned on the same side, allowing both
tentacles to be streamed in the same direction.

The animal was dislodged from the alga and
was observed to swim feebly by a series of
undulating wave movements of the thin peri-
pheral region. It was also observed to gain the
surface film of water in a shallow dish and to
float inverted on the surface film by completely
expanding its under-surface. While floating in
this manner the tentacles were fully streamed
several times. On one occasion the tentacles
touched the bottom of the dish and appeared
to adhere to it for a short period. During this
time they were slowly contracted, pulling the
animal along.

When stimulated with a needle, when either
crawling or floating, the tentacles were retracted
very rapidly and completely. After a short

period, either one or both were very slowly
expanded in stages, each stage being inter-
spersed with further rapid complete contrac-
tions. No feeding activity was observed.

IDENTIFICATION

Platyctenean ctenophorans have been re-
corded from many parts of the world, from
Greenland (Mortensen, 1912) to Antarctica
(P. Dayton and G. A. Robilliard 1968 in
Gordon 1969), with records from practically
every contient. However, they are still such
rare and unusual animals that they have promp-
ted study whenever they have been discoverd
and many new taxa have resulted. This has
been especially evident when a specimen has
been discovered for the first time in a marine
faunal zone or geographical region in which
this group of animals has not previously been
recorded. This phenomenon, coupled with the
absence of any clearly defined taxonomically
useful characters, has led to a proliferation of
specific and generic names and taxonomic
confusion.

Several partial revisions of the group, have
been undertaken, principally by Dawydoff
(1936, 1938), Komai (1934) and others,
while a good general account is provided by
Hymen (1940). Following this latter work the
present specimens are referred to the genus
Coeloplana because of the presence of erectile
dorsal papillae, the absence of comb-plates, and
because the statocyst and tentacles are in hidden
tentacular sheaths. The specimens are tenta-
tively identified as Coeloplana willeyi Abbott
(1907), following Matthews and Townsley
(1964) and Gordon (1969), because the size,
colour and number of dorsal papillae fall within
the variation range of this species. However it
is recorded as such, more to provide a con-
venient label for future reference than with
any idea of taxonomic exactitude.

Acknowledgements

We would like to thank Mr Phil Hollis for
first discovering these animals and making
them available for study.

References

Abbott, J. F., 1907. The morphology of Coeloplana.
ZooL Jl. (Anat. Ontog.) 24: 41-70.

CREEPING CTENOPHORAN

Figs. 1, 2 and 3 — Series of drawings of Coeloplana
willeyi Abbott of the living animal showing
the general body shape and the tentacles.

Fig. 4 — Central statocyst.

Fig. 5 — Animal floating on surface film, using the
tentacles for locomotion.

BRIAN J. SMITH and RHYLLIS J. PLANT

Dawydoff, C, 1936. Les Ctenoplanidae des eaux de
l'lndochine Francaise. Etude systematique. Bull,
biol. Fr. Belg., 70: 456-486.

, 1938. Les Coeloplanides Indochinaise.

Arch. tool. exp. gen. 80: 125-162.

Gordon, D. P., 1969. A Platyctenean Ctenophore
from New Zealand. N.Z. 31. mar. Freshwat. Res.,
3: 466-471.

Hyman, L. H., 1940. The Invertebrates. Part 1. Pro-
tozoa through Ctenophora. McGraw-Hill, New
York.

Komai, T., 1934. On the structure of Ctenoplana.
Mem. Coll. Sci. Kyoto Univ., Ser. B. 9: 245-256.

Matthews, D. C. and S. J. Townsley, 1964. Addi-
tional records of Hawaiian Platyctenea (Cteno-
phora). Pacif. Set., 18: 349-451.

Mortensen, T., 1912. Ctenophore I. Tjalfiella tris-
toma Mrtas. Dan. Ingolf. Exped., 5(2): 3-59.

Stephenson, T. A. et al., 1931. The structure and
Ecology of Low Isles and other reefs. Sclent.
Rep. Gt. Barrier Reef Exped., 3: 17-112.

SYMBIOCLADIUS AURIFODINAE sp. nov. (DIPTERA, CHIRONOMIDAE),
A PARASITE OF NYMPHS OF AUSTRALIAN LEPTOPHLEBIIDAE

(EPHEMEROPTERA)

By H. B. N. Hynes

Department of Biology, University of Waterloo, Ontario, Canada

Summary

Symbiocladius aurifodinae sp. nov., an orthoclad chironomid parastic on nymphs of the
mayfly genus Atalophlebioides, is described from mountain streams in Victoria. Descriptions
are based on mature pupae and larvae collected during the summer. It is concluded that there
is probably only one generation per year. This is the first record of Symbiocladius from
Australia, and it is shown that this species is closely related to S. wygodzinskvi Roback from
Argentina.

Introduction

Fontaine (1964) and Arvy and Peters
(1973) have reviewed the literature on the
larvae of Chironomidae that are found in asso-
ciation with mayfly nymphs. It appears that
the only species which actually feed on the
tissues of their hosts are members of the genus
Symbiocladius, and most records are from
the flattened nympths of the Heptageniidae of
the Northern Hemisphere. There is, however,
one record of larvae on a species of Lepto-
phlebiidae from North America (Mayo 1969),
and careful description of another type of
larva from a flattened leptophlebiid from sou-
thern South America (Roback 1965). It may
be noted here that flattened leptophlebiid
nymphs replace the Heptageniidae in the
Southern Hemisphere.

The larvae found by Mayo (1969) on
nymphs of Thraulodes were identified as Sym-
biocladius, but it is clear from her description
and figures that they differ considerably from
the other described species. The larvae, for
instance, seem not to be parasitic, they have
well-formed head capsules and they retain
eyespots, caudal bristles and anal gills. They
seem, in fact, to be not unlike the phoretic
genus Plecoteracoluthus, which occurs on per-
lid stoneflies (Steffan 1965) and Megaloptera
(Hilsenhoff 1968).

The specimens from Argentina, on the other
hand, are clearly parasitic and damaging to
their hosts, which were tentatively identified
as Thraulodes. It was intriguing therefore,
especially in view of the similarity of the biotas

of southern South America and Australia, to
find a very close relative on nymphs of the
leptophlebiid Atalophlebioides in streams on
the Great Dividing Range in Victoria.
THE MATERIAL
The specimens were obtained during monthly
collections (June 1971 to June 1972) of the
fauna of several streams that were used in the
study of the life histories of stoneflies. The
methods used are described by Hynes and
Hynes (1975), where more information on the
streams is given. Mayfly nymphs carrying larvae
or pupae of Chironomidae were found in only
three of the 11 stream stations that were in-
tensively studied.
These were:

Crown Creek above Woods Point (map re-
ference 424367), 2300 ft, a cool (max.
16i°C), swift stream 5-10 m wide and up
to 40 cm deep, with a stable bed of rocks
and shingle.

Godfreys Creek below Frenchman's Gap
(421374), 2500 ft, a cool (max. 13*°C),
fairly swift shallow stream 2-3 m wide, with
a stable stony and gravelly bed containing
some silt.

Delatite River below Sawmill Settlement

(434423), 1900 ft, a cool (max. 16*°C),

swift, turbulent river about 15 m wide and

up to at least 1 m deep, with a stable bed

of boulders, stones and coarse sand.

No specimens of Atalophlebioides, which is

a common genus in stony streams, were seen

with chironomid larvae in any of the many

other streams that we visited in Victoria. How-

H. B. N. HYNES

ever, one pupa was taken, and unfortunately
lost in an attempt to breed it out, in Leather-
barrel Creek, N.S.W. (615491) on January 8,
1974.

Two larvae were first noticed in the collec-
tion made on October 26, 1971, in Delatite
River, one was found on November 24 in
Crown Creek, and several were found in God-
freys Creek on December 28. They persisted
in small numbers until February and March in
the two creeks; in December there were many
pupae, and in January and February only pupae
(one pre-pupa in March) were obtained. In
Delatite River only pupae were found on De-
cember 28 and no specimens were collected on
or after January 25. These findings possibly
indicate only a single generation per season
for the chironomid. It seems that at least some
species of the host mayfly, which is present at
all times, are uni voltine ( Duncan 1 972 ) .
Thirty-seven larvae and pupae were collected
in total, 20 from Godfreys Creek and 12 from
Delatite River, and four mayflies carried empty
shrouds from which, presumably, pupae had
emerged.

The Hosts

The host mayflies appear to be all of the
same species of Atalophlebioides (Figure 1),
which it is possible may later become specifi-
cally identifiable by its comb-like tarsal claws
and by the peculiarly thickened distal margins
of two of the segments near the bases of the
three tails (Figures 1, C and D). No counts
of uninfected specimens were made, but it is
estimated that only about 1% carried the
chironomid.

Infected nymphs are all middle-sized, 3i to
61 mm long, whereas fully developed nymphs
were 8-8i mm long. There was good evidence
that the chironomid caused stunting rather
than that smaller nymphs were selected for
infestation. The larvae were attached laterally
to the thorax, indifferently as to side: 18 out
of 41 were on the right. When wing pads were
present the one nearest to the parasite was
always reduced (Figures 1, A and B) and the
entire development of the nymph seems to have
been retarded. For instance, the uninfected
nymph of which the mesonotum is shown in

Figure 1, F, was the same size and, as judged
by the developing male eyes, in the same instar
(probably the penultimate) as the specimen
shown in Figure 1, B. Also, the most fully
developed infected nymph, a 6i mm male that
still carried an empty pupal shroud, seemed to
be physiologically near emergence in that its
better developed wing pad enclosed a folded
structure; but its wings were tiny and distorted
(Figure 1, E) as compared with normal speci-
mens in late instars (Figure 1, G and H). It
may also be significant that the only chironomid
collected on March 29 was a pre-pupa 1*75
mm long, which is shorter than the smallest
formed pupa that was collected, and it was
taken on one of the two smallest infected
mayflies (3i mm). It was also the only larva
collected after the end of December. One may
suppose that it infested a nymph that was too
small or too unhealthy for ordinary develop-
ment to occur.

The parasite larvae were completely enclosed
in a tough membranous shroud, as has been
described for other species of Symbiocladius
(Codreanu, 1939; Roback, 1965), and they
lay alongside the nota, sometimes with the tail
tucked under the wing pad as shown in Figure
1, A. The larval head was directed forward
or backward, apparently indifferently (13 out
of 25 were forward), but I could find no
trace of lesions on the nymphs. It seems clear,
however, that, as with the European species
(Codreanu, 1939), the host must supply the
food.

The pupae were all attached as shown in
Figure 1, B, with their heads over the mayfly
abdomen, and it appears from the empty
shrouds that they leave by a dorsal rip. There
was no relationship between the sex of the
host and that of the pupal parasites; all four
possible combinations were found among the
12 pupae collected.

THE PARASITE

Six specimens, two larvae and four pupae,
were used for the preparation of microscope
slides for comparison with the clearly closely
similar Argentinian species S. wygodzinskyi
(Roback, 1965). The mountant was Euparal.

LEPTOPHLEBIIDAE NYMPHS PARASITES

Fig. 1 — A and B, Nymphs of Atalophlebioides with
a larva and pupa of Symbiocladius aurifo-
dinae (larger scale line 1 mm) : C and D,
claw and base of filum terminale of Atalo-
phlebioides (upper scale line 0-1 mm): E-H,
mesonota of Atalophlebioides (larger scale
line 1 mm), E, an infected specimen 6i mm
long, F-H, uninfected specimens 6i, 7i and
8i mm long. H, from Delatite River Decem-
ber 28, 1971, E, Godfreys Cr. February 25,
1972, rest from Godfreys Cr. December 28,
1971.

Genus Symbiocladius Kieffer
Subgenus Acletius Roback
1965 Entomol. News 76: 114-115.

The Australian material agrees in most
respects with Roback's definition of Acletius.
These include the haired eyes of the adult
(Figure 2, E and H), the subequal tibial spurs

(Figure 2, D), the long pectinate empodium, the
three long basal spines on the claws (Figure
2, G), the latero-dorsal position of the larvae
on the host (Figure 1, A), the five lateral teeth
on the labial plate (Figure 3, B and E) and
the two teeth on the mandibles (Figure 3, C).
There are, however, small differences. The
antennae of the female have seven segments,
rather than six (Figure 2, H), and the two
mandibular teeth of the larva are subequal
rather than being one robust and one accessory
(Figure 3, C). I was unable to observe the
palpal segments of the adult.

Symbiocladius (Acletius) aurifodinae sp. nov.

Male described from pupal material; 2-2
mm. Head and thorax dark brown, abdomen

H. B. N. HYNES

Fi £- 2 — Sytnbiocladius aurifodinae. A and B, D-G
holotype, H and I, allotype. A, genitalia and
part of pupal skin (smaller scale line 0-1
mm): B, tip of dististyle (larger scale 01
mm): C, ditto of specimen from Godfreys
Cr December 28, 1971: D, tibial spurs of
hind leg (smaller scale line 01 mm): E,
optical section of eye margin (larger scale
line 0-1 mm): F, pupal thoracic respiratory
organ (smaller scale line 01 mm); G, claw
of hmd leg (smaller scale line 001 mm): H
and I, head and abdominal tip of female
(smaller scale line 01 mm).

brown with paler patches at bases of hairs.
Antennae with at least 14 segments; eyes hairy,
the hairs about as long as the diameter of the
facets (Figure 2, E). Thorax apparently with
bristles omly on scutellum. Pronotum narrow
and collar-like. Legs with subequal tibial spurs
and with claws with three basal hairs (Figure 2,
D and G); segment ratios as in Table 1.
Genitalia as Figure 2, A and B, but note that
the number of bristles at the tip of the dististyle
is three in the type and five in the other speci-
men (Figure 2, C).

TABLE 1

Symbiocladius aurifodinae ratios of leg seg-
ments of type specimens (100 = 07 mm)

tarsal segments
femur tibia 12 3 4 5

Male

fore 100 200 130 88 49 16 17
mid 99 146 131 34 27 16 15

hind

134 163 108 60 33 21 21

Female
fore

161

225 171 64 37 29 29

Female described from pupal material; 31
mm. Similar to male, and also to S. wygod-
zinskyi Roback (Figure 2, H and I). It has,
however, seven antennal segments, no ventral
hair on the antennal pedicel, and it lacks the
paler spots at the bases of the abdominal
hairs.

Pupa, length, male 2-2 to 2-8 mm (holo-
type 2-2), female 2-3 to 3-8 (allotype 31);
small respiratory trumpets present (Figure 2,

LEPTOPHLEBIIDAE NYMPHS PARASITES

Fig. 3 — Symbiocladius aurifodinae, larval appendages
(scale line 001 mm): A, antenna: B, labial
plate: C. mandible: D, Labrum: E, labial
plate and tip of labrum: F, maxillary palp:
A-D, head capsule of holotype: E and F,
head capsule from a female pupa from
Delatite R. December 28, 1971.

F) . Cuticle thin and pale brown, without spines.
In the male a small featureless tail fin (Figure
2, A); genital sacs elongate (Figure 2, A,
where the tips of the sacs are missing). Note
that Roback (1965) states that in S. wygod-
zinskyi the anal fins are twice the length of
the genital sacs, but this does not agree with
his figure. In mature female pupae, in which
the abdomen is distended with eggs, the anal
fin is hardly apparent.

Larva, the larvae range in length from 0-5
to 2-1 mm (two that are certainly last instar,
with developing appendages, are 2 and 2-1
mm). Head pale yellow with a darkened hind
edge; capsule widely open behind, almost a
hemisphere. No eyespots. Body light brown
with well formed anterior prolegs and small
posterior ones; no posterior hairs or gills. An-

tennae small and little sclerotized; number of
segments uncertain (Figure 3, A). Labial plate
broad with five lightly sclerotized teeth on each
side (Figures 3, B and E). Maxilla shows only
a small lightly sclerotized ring with several
central conical structures (Figure 3, F). Man-
dibles conspicuous hooks with two accessory
teeth (Figure 3, C). Labrum with an apical
swelling bearing about five lightly sclerotized
teeth (Figures 3, D and E).

MATERIAL EXAMINED

Descriptions made mostly from the six
mounted specimens.

Holotype — mature male pupa, Crown Cr.,
Jan. 27, 1972.

Allotype — mature female pupa, Godfreys Cr.,
Jan. 27, 1972.

Paratypes — male pupa and 2 last instar larvae,
same data as allotype.
— head capsule of larva extracted
from case of female pupa, Dela-
tite R., Dec. 28, 1971.

H. B. N. HYNES

Other Material

— 2 larvae, 1 prepupa, 1 female pupa,
Crown Cr., Nov. 24 and Dec. 28,
1971, and March 29, 1972.
— 11 larvae, 2 male pupae, 2 female
pupae, Godfreys Cr., Dec. 28,
1971, and Feb. 25, 1972.
— 9 larvae, 3 female pupae, Delatite
R., Oct. 26, Nov. 23 and Dec. 28,
1971.
All the specimens were collected by H. B. N.
and M. E. Hynes. The types, paratypes and
much of the other material has been presented
to the National Museum of Victoria in Mel-
bourne. Two larvae and one pupa from Delatite
R., and four larvae and two pupae from God-
freys Cr. remain in the author's collection. The
species is named for the goldmines which are
still active around Woods Point near to which
most of the specimens were collected.

Although in the absence of fully developed
specimens, it is difficult to be certain, adults
would probably run down to the genus Cric-
topus in Freeman's (1961) key to the Aus-
tralian Orthocladiinac. The larvae and pupae
are, however, very different from those of CricCh
topus. They are also quite distinct from those
of Trissocladius, with which the genus Symbio-
cladius has sometimes been combined, as was
already pointed out by Saether (1969).

S. aurifodinae is clearly very closely related
to S. wygodzinskyi, but there are a few dif-
ferences. The male has more antennal flagellar
segments, at least 14 as opposed to 13, and the
bristles at the dististyle tip are pointed not ovate.
Th|C female has one more antennal segment.
The pupa lacks dorsal spines and possesses
a small respiratory organ. The larva has, ap-
parently, much less robust lateral labial teeth
and a wider central part to the labial plate,
and its second mandibular tooth is larger. Also
the tip of the labium which carries the spines
is more swollen, and there seem to be no lateral

spines below it such as are figured for S. wygo-
dzinskyi by Roback. These seem to be very
small changes after what must be many tens of
millions of years of isolation of the two con-
tinents.

Acknowledgements

I have previously acknowledged the help
from many people and organizations that al-
lowed me so rewarding a sabbatical year in
Australia. My principal benefactors were the
University of Waterloo, Monash University,
and the National Research Council of Canada.
I remain in their debt.

References

Arvy, L. and W. L. Peters, 1973. Phoresies, bio-
coenoses et thnnatocoenoses chez les ephemerop-
tcres. Proceedings of the First International Con-
ference on Ephemeroptera 1970, Tallahassee,
Florida, 244-312.

Codreanu, R., 1939. Recherches biologiques sur un
chironomid Symhiocladius rhithrogenac (Zavr.)
ectoparasite 'cancerigene' des ephemeres torren-
ticoles. Arch. ZooL exp. gen. 81: 1-283.

Duncan, M. J., 1972. The life histories of Ephemerop-
tera from two Victorian streams. B.Sc. Honours
thesis. Department of Zoology, Monash Univer-
sity.

Fontaine, J. 1964. Commensalisme et parasitisme
chez les larves d'ephemeropteres. Bull, mens. Soc.
Lyon, 33: 163-174.

Freeman, P., 1961. The Chironomidae (Diptera) of
Australia. Aust. J. Zool. 9: 611-737.

Hilsenhoff, W. L., 1968. Phoresy by Plecoptera-
coluthus downesi on larvae of Nigronia serri-
corm's. Ann. Entomol. Soc. Amer. 61: 1622-23.

Hynes, H. B. N. and M. E. Hynes, 1975. The life
histories of many of the stoneflies (Plecoptera)
of southeastern mainland Australia. Aust. J. mar.
Freshwat. Res. 26: 113-53.

Mayo, V. K., 1969. Nymph of Thraulodes speciosus
Traver with notes on a symbiotic chironomid.
Pan-Pacif. Ent. 45: 103-112.

Roback, S. S., 1965. A new subgenus and species of
Svmbiocladius from South America. (Diptera:
fendipedidae). Entomol. News 76: 113-122.

Saether, O. A., 1969. Some nearctic Podonominae,
Diamesinae, and Orthocladiinae (Diptera, Chiro-
nomidae). Bull. Fish. Res. Bd. Can. 170: 1-154.

Steffan, A. W., 1 965 . Plecopteracoluthus downesi
gen. et sp. nov. (Diptera: Chironomidae), a
species whose larvae live phoretically on larvae
of Plecoptera. Canad. Ent. 97: 1323-44.

THE ASCIDIAN FAUNA OF WESTERN PORT, VICTORIA, AND A
COMPARISON WITH THAT OF PORT PHILLIP BAY

By Patricia Kott

Queensland Museum
Abstract

The taxonomy of 59 species of ascidians from Western Port and Port Phillip Bay,
Victoria, is discussed. The ascidian fauna of Western Port is markedly more diverse than
that of Port Phillip Bay. The biogeographical affinities of the species are assessed and the
implications of the differences in species composition in the two areas are investigated.

Introduction

A previous collection of ascidians from Port
Phillip Bay has been reported on by Millar
(1966) but prior to that no major work has
been devoted to the ascidian fauna of Victoria.
The greater part of the present material has
been collected for the National Museum of
Victoria by the Underwater Research Group
(Western Port Survey). Additional records of
species occurring in Western Port, available
from independent collections made by Mrs. J.
Watson and Mr. K. Duke on parts of the
adjacent Victorian coast, from Mallaooota
near the Victorian — N.S.W. border to Portland
Harbour and Cape Nelson, have been included
in the present work.

These collections are of particular interest
in relation to the fauna of St. Vincent Gulf
where large collections have recently been
made and reported on (see Kott, 1972 a, b).
Information on the better known fauna of Port
Jackson and Moreton Bay to the north is also
available. (Kott, 1952; 1957; 1962; 1963;
1972 c, d). These locations are all large em-
bayments in the Australian coastline, essen-
tially marine, and tidal. They are, however,
all protected from the direct swell of the
southern ocean, and receive some fresh-water
runoff from the water-ways emptying into them
and from the shores surrounding them.

The distribution of this sessile ascidian fauna
is limited by the short free-swimming life of
the pelagic larvae. Consequently species adap-
ted to protected localities could be restricted
in their distribution by lack of suitable sites
for settlement on the open coast. The phylo-
genetic relationships of the ascidian fauna of

each of these sheltered embayments are there-
fore of special zoogeographic and ecological
interest in view of the likelihood of isolation
of endemic and relict species.

There are 59 species in the collections, of
which one, Ciona intestinalis is probably in-
troduced. These species, are set out in Tables
1 and 2 together with others previously re-
corded from Port Phillip Bay and Western
Port but not represented in these collection.
Aspects concerning the biogeography and habi-
tat of the ascidian fauna of these locations is
discussed below.

Species List

APLOUSOBRANCH1A
CIONIDAE

Ciona intestinalis
CLAVELINIDAE

Clavelininae

Oxycorynia pseudobaudinensis n. sp.

Podoclavella cylindrica

Holozoinae

Atapozoa mirabilis

Sycozoa pedunculata

Sycozoa cerebriformis
POLYCITORIDAE

Eudistoma pyriforme
POLYCLINIDAE

EUHERDMAN1INAE

Pseudodistoma cereum
Dumus arenijerus

POLYCLININAE

Polyclinum marsupiale
Aplidium depressum
Aplidium lobatum
Aplidium triggiensis

PATRICIA KOTT

Synoicum hypurgon

Synoicum sp.?

Sidneyoides tamaramae
DIDEMNJDAE

Trididemnum cerebrijorme

Trididemnum cyclops

Didemnum candidum

Didemnum spongioides

Didemnum skeati

Didemnum moseleyi

Didemnum patulum

Didemnum turritum

Didemnum augusti

Didemnum roberti

Didemnum lambitum

Lissoclinum fragile

Lissoclinum ostrearium

Diplosoma translucida

Diplosma rayneri

Polysyrwraton orbiculum

Polysyncraton victoriensis n. sp.
PHLEBOBRANCHIA
ASCIDIIDAE

ASCIDIINAE

Phallusia depressiuscula
Ascidia Sydney ensis
Ascidia gemmata
STOLIDOBRANCHIA
STYELIDAE

BOTRYLLINAE

Botrylloides leachii
Botrylloides nigrum

POLYZOINAE

Symplegma viride
Amphicarpa diptycha
Polyandrocarpa lapidosa

STYEL1NAE

Polycarpa thelypanes
Cnemidocarpa etheridgii
PYURIDAE
Pyura australis
Pyura cataphracta
Pyura irregularis
Pyura albanyensis
Pyura lepidoderma
Pyura scores biensis
Pyura stolonifera praeputialis
Halocynthia hispida
Herdmania momus

Microcosmus australis

Microcosmus nichollsi

Microcosmus helleri

Microcosmus stolonifera

Microcosmus squamiger

MOLGULIDAE

Molgula mollis

Molgula sabulosa

SYSTEMATICS

Ciona intestinalis Linnaeus

Ciona intestinalis Linneaus, 1767, p. 1087. Kott, 1952,
p. 319 for synonymy and description.

New Records: Port Phillip Bay (Oil wharf,

Yarra River; artificial reef).

Distribution: See Kott, 1952.

Remarks: Kott (1969) has suggested that the

cosmopolitan occurrence of this species, which

is recorded from harbours and wharf piles in

all regions outside the Antartic, is due to its

transport on ships' hulls.

Oxycotynia pseudobaudiensis sp. nov.

(Fig. 1)

Clavelina baudinensis Kott, 1957, p. 87 (part: speci-
men with larger larvae) ?Millar, 1966, p. 363. Kott,
1972a, p. 4.

Type Locality: Laverton Bay (Victoria)
Holotype: Australian Museum A.M. Y1113.
Paratypes: W. Aust. Rottnest I., AM Y1112
(Kott 1957). S. Aust.: Carickalinga Head,
South Australian Museum S.A.M. E876;
Rapid Head S.A.M. E 877 (Kott 1972a).
Vict. (Western Port): Balnarring Beach, A.M.
Y1122 (Kott 1957); Crawfish Rock; Flinders
Jetty, N.M.V. (new records).
Description: The colonies are 5-8 cm high
and the wider terminal part of the head is 2
cm in diameter terminally. In the upper half
of the colonies the test is delicate and some-
times glassy and transparent and encloses the
body of the zooids which are never separate.
On the upper surface the test forms only
slight rounded protruberances over the anter-
ior aspect of the zooids. In some colonies the
zooid bearing upper part may be subdivided
into several lobes. The slightly bulbous stalk
narrows toward the base and is only slightly
longer than half the height of the colony. The
test of the stalk is firm and opaque and some-
times slightly leathery externally.

ASCIDIAN FAUNA OF WESTERN PORT

Zooids are from six to eight mm. In pre-
servative they are a bluish colour and have
dark accumulations of pigment in the mid line
dorsal and ventral to the branchial
siphon. There are 12 to 20 longitudinal
muscles on the thorax and, depending on their
degree of coalescence, they may vary in num-
ber on each side of the body. From six to eight
of the most ventral bands are aligned at a
slight angle with the longitudinal axis of the
body and break up into branches over the
anterior half of the endostyle. Of the remain-
ing longitudinal muscle bands more than half
extend into the branchial siphon and the others
into the atrial siphon. Posteriorly the bands
extend along both sides of the abdomen. There
are from 16 to 20 rows each of 20 to 30
rectangular stigmata with a well developed
transverse membrane between each row. In
the mid dorsal line this membrane is expanded
into the usual triangular, pointed languets.

The gut forms a simple and fairly short loop
(seldom longer than the thorax), enclosing
the gonads. The anus opens at the base of the
peribranchial cavity and is bordered with min-
ute rounded lobes. The stomach has no true
structural folds. Tt is present in the middle to
posterior one third of the abdomen. There is
no prestomach.

Larvae are present only in the colonies from
Rottnest Island and Laverton Bay (see Kott,
1957: larger larvae). They are large, 0-9 mm
long with the tail wound threequarters of the
way around the body. Triradiate papillae are
supported around a flattened frontal plate.
The adhesive cells rise in a cone from tho
centre of a depressed area which forms a
fairly primitive papillary sucker or cup. The
embryos appear to start their development in
the proximal part of the oviduct and complete
it in the right side of the peribranchial cavity
where they demonstrate a wide range in stages
of development. The most mature embryos
are present anteriorly.

Remarks: The separate identity of the present
species was first suggested by the different
larvae present in colonies from Rottnest and
Victoria which had all been assigned to the

species C. baudinensis Kott, 1952. Most of the

previously described specimens of that species
excepting only those recorded by Millar (1966)
have been re-examined.

Clavelina baudinensis from Cape Vlamingh,
Rottnest Island and from Laverton Bay have
small larvae (0-5 mm) in which the simple
papillae without accessory suckers are sup-
ported around the anterior end of the body
which is not separated into a frontal plate.
In these larvae the tail completely circles the
body. These colonies can. be distinguished
from the present species mainly by their long-
er, narrow and cylindrical stalk. The anterior
extremity of the zooids project more from the
anterior surface of the test than in O. pseudo-
baudinensis. Zooids of C. baudinensis exam-
ined have a maximum of 12 longitudinal thor-
acic muscles of which only a single band
subdivides across the mid-line, ventral to the
branchial aperture. In C. baudinensis there
appears to be a more restricted range in the
stage of development of embryos in the peri-
branchial cavity. Although some eggs are
present in the oviduct they do not appear to
start their development there as in the genus
Pycnodavella. In its colony and zooid form,
C. baudinensis does appear to be closely re-
lated to O. pseudobaudinensis; however, its
larvae and the degree to which eggs are ap-
parently fertilised in the atrial cavity suggest
that it is a more primitive species.

In O. pseudobaudinensis the oblique arrange-
ment of the ventral thoracic muscles effects a
depression of the anterior part of the thorax
and draws it towards the postero-dorsal part
of the thorax. The atrial aperture simultan-
eously becomes terminal as in P. cylindrica,
thus facilitating the liberation of large larvae.
In C. baudinensis larvae are smaller and more
easily liberated through the normally oriented
aperture; and the more parallel arrangement of
longitudinal thoracic muscles does not apocar
to affect the relative position of the siphons.
In neither of these species is the whole ventral
surface withdrawn toward the postero-dorsal
part of the thorax as is the case in Podocla-
vella cylindrica where there is a special brood
pouch ensuring the retention of embryos.

PATRICIA KOTT

Podoclavella cylindrica (Quoy and Gaimard)

Polyclinum cylindrica Quoy and Gaimard, 1834, p.
618.

Podoclavella cylindrica; Kott, 1972a, p. 5 and syno-
nymy; 1972b, p. 167.

New Records; Western Port (Flinders Jetty,
Balnarring Beach). Ram Head (18 mis south
of Mallacoota Inlet, 6 m).
Distribution: W. Aust: Rottnest Island, Fre-
mantle, Albany; S. Aust.: St. Vincent Gulf,
West Island, Wright Island; Vict.: Bass Strait,
Port Phillip Bay. The greatest recorded depth
for the species is 22 metres (West Island,
Kott, 1 972 ) . The species is known from
sheltered caves and under ledges.
Description: The present colonies generally
have a firm basal common stalk and zooids
are supported in the less firm terminal test
and are independent of one another anteriorly.
The usual blue pigment spots are present an-
teriorly and thoracic musculature extends
obliquely from the ventral border to the pos-
tero-dorsal aspect of the body. Only in speci-
mens from Western Port Bay are the zooids
arranged around a central stalk which is 24
cm long and 2-4 cm in diameter, thickest
terminally where it breaks into branches.
Larvae of the usual form without ampullary
lobes, are present in the brood pouches of this
colony.

Remarks: In P. cylindrica contraction of the
oblique thoracic musculature causes the fore-
shortening of the dorsal length of the thorax by
drawing the postero-dorsal corner ventrally and
anteriorly. The branchial aperture is simultan-
eously withdrawn leaving the atrial aperture
terminal and anal and gonadial openings ad-
jacent to it. The developing embryos are re-
tained however, in a pouch from the dorsal
surface, thus avoiding early liberation which
could otherwise result from contraction of the
thorax.

The relationship of specimens in which
there is a central stalk to those that are sup-
ported upright and parallel to one another on
a basal membrane is not known. Another
specimen of the former type in the collection
of the National Museum of Victoria is 60 cm
long and resembles Distaplia cylindrica from

the Antarctic (see Kott, 1969). The zooids
in both forms are similar in every respect and
the colonies appear to represent the same
species. It is possible that the long axial
stemmed forms are from deeper water.

Atapazoa mirabilis Kott
(Fig. 2)

Atapazoa mirabilis Kott, 1972b, p. 168.
New Records; Western Port (Tankerton jetty)
Distribution: The species has previously been
recorded from S. Aust. (Elliston Bay).
Description: A single colony only is available.
It is massive and irregular, 14 cm long and
6 cm wide. It is composed of fairly thin layers
of zooid bearing test that coalesce so that the
colony is traversed by spaces. The atrial siphon
from the postero-dorsal corner of the thorax
is characteristically long and posteriorly direc-
ted. Both the branchial and atrial apertures
are bordered by six distinct lobes. There are
three rows each of about 12 stigmata in each
row. The horizontal gut-loop consists of a
fairly long oesophagus, rounded stomach and
wide intestinal loop. There is a single large
ovum attached to the zooid from a region in
the loop of the gut. The position of the brood
pouch is apparently abdominal, rather than
thoracic, and is reminescent of the situation in
the Didemnidae.

Remarks: The species has been recorded pre-
viously only from Elliston Bay in South
Australia. It is possible that it is endemic to
the southern coast although the type location,
on the floor of a cave, is only accessible to
collectors equipped with SCUBA.

Sycozoa pedunculata (Quoy and Gaimard)

Aplidie pedunculatum Quoy and Gaimard, 1834, p.
626.

Sycozoa penduculata; Kott, 1972c, p. 234 and syno-
nymy.

New Records: Western Port (Rutherford
Channel); Port Phillip Bay (no location, arti-
ficial reef). South-east Portland.
Distribution: W. Aust.: Cockburn Sound, King
Georges Sound; Tas.: d'Entrecastaeux Chan-
nel, Derwent Estuary, Furneaux Group; S.
Aust.: St. Vincent Gulf; Vict.: Western Port,

ASCIDIAN FAUNA OF WESTERN PORT

Port Phillip Bay, Lakes Entrance; Qd. : More-
ton Bay.

Discription: The usual large inverted conical
heads on long slender stalks with basal tufts
of roots. There are deep V-shaped furrows
between each double row of zooids and the
branchial apertures from each row of zooids
open into each side wall of these furrows. The
branchial openings are thus protected to some
extent and the furrow provides an immediate
microenvironment outside the openings. The
apex of a rounded ridge between two of these
furrows lies over the common cloacal canal.
There are very large common cloacal openings
around the outside of the flattened free ends
of the lobes.

Larvae are present in colonies from South-
east Portland. They have the usual anterior
papillae, an otolith, but no ocellus, and a
short broad tail extending only three quarters
of the ways around the body of the larva.

Sycozoa cerebriformis (Quoy and Gaimard)

Aplidie cerebriforma Quoy and Gaimard, 1834, p.

625.
Sycozoa cerebriformis; Kott, 1972a, p. 8 and syno-
nymy.

New Records: Western Port (Crawfish Rock,
Tankerton Jetty). Portland Harbour (5-10 in
on rocks forming jetty).

Distribution: N.W. Aust; S. Aust.: St. Vincent
Gulf; Vict. : Balnarring Beach (Western
Port), Port Phillip Bay; N.S.W.: Jervis Bay,
Port Stephens, Port Hacking, Port Jackson.
South Africa. It has been recorded from
5-40 m.

Description: Specimens are sturdy fan-shaped
colonies with short stalks. The fan or zooid
bearing portion may extend into a thick
undulating lamellae. Common cloacal aper-
tures are present along either side of the flat-
tened free edge of the lamella. The double rows
of zooids converge from this outer edge of the
lamella down toward the top of the stalk.
Remarks: The specimens from deeper water
at Crawfish Rock are larger than the colony
taken in shallower water. It has already been
observed (Kott, 1972a) that the species fav-
ours areas where there are steady but not
strong unidirectional currents, and no surge

or turbulence. The fan-like colony shape is
apparently adapted to take maximum advan-
tage of this type of environment. This species
appears to be confined to more sheltered
locations than S. pedunculata, very often where
there is some turbidity and a muddy substrate
although in the absence of a larval ocellus
the light conditions are not likely to affect its
settlement.

It has been recorded from a wider circum-
polar range than S\ pedunculata but although
it has been taken from north western Australia
it is not recorded from Moreton Bay and it
has not been recorded from Tasmania. Its
latitudinal range is therefore more limited. It
has not been taken from depths greater than
40 metres. Its distribution suggests that it
may represent a relict species confined within
embayments in relatively shallow water where
it can best take advantage of sheltered con-
ditions.

Polycitor giganteum (Herdman)

Polyclinum giganteum Herdman, 1899, p. 79.
Polycitor giganteum; Kott, 1972a, p. 9 and synonymy.

New Records: Western Port (Crawfish Rock,
Tankerton Jetty). Ram Head (18 mis south
of Mallacoota Inlet, 6 m).
Distribution: A wide circum-Australian dis-
tribution from Rottnest Island (W.A.) and
across the southern coast to Port Jackson (N.
S. W.).

Description: One specimen from Crawfish
Rock is more or less flattened and sessile,
about 13 cm in diameter but only 5 cm high;
another specimen has the usual rounded gel-
atinous head narrowing to a waist before ex-
panding into a wide sandy base, possibly
embedded in the substrate. The test is char-
acteristically transparent and gelatinous and
the zooids are large, radiating out from the
base of the colony to open by separate
apertures around the head.
Remarks: The species appears to favour rocky
substrates where -a firm adhesion can be
effected, thus satisfying the requirements of
a large, inflexible colony that is fixed by only
a small area of the base. The species is found
equally in embayments and on the open coast.

PATRICIA KOTT

Eudistoma pyriforme (Hcrdman)

(Fig. 3)

Psammaplidium pyriforme Herdman, 1886, p. 419.
Eudistoma pyriforme; Kott, 1972a, p. 9 and synonymy.

New Records: Western Port (Crawfish Rock).
Ram Head (18 mis. south of Mallacoota Inlet,
6 m).

Distribution: Palao and Gilbert Islands. Qd.:
The Great Barrier Reef, in the Pacific; S.
Aust.: St. Vincent Gulf. Madagascar.
Description: The colony from Mallacoota Inlet
is flattened and firm with sand absent only
from the surface layer of test which has
brownish-purple spherical pigment cells. The
colony from Crawfish Rock is irregular and
investing, with a dense sand inclusion through-
out the test, making it rather hard and ob-
scuring the arrangement of the zooids. There
are about 10 fairly wide muscle bands down
either side of the thorax and an almost con-
tinous layer of circular muscles. The circular
muscles on the siphons are well developed but
do not form definite sphincters. The oesopha-
gus is of medium length opening into a large
shield shaped and smooth stomach halfway
down the abdomen; the duodenal area is long,
and in a contracted abdomen is bent in an
S-shape. The intestine bends anteriorly after
leaving a spherical posterior stomach, and
forms a loop opposite the duodenum when the
abdomen is contracted. The rectum extends
anteriorly, to the peribranchial cavity, and is
straight.

Remarks: Although Hastings (1931) regarded
the loop in the gut as diagnostic of this
species it has been observed in other species
where the abdomen is contracted. There are few
reliable diagnostic characters available in this
genus where colony shape is variable, no
systems are formed, and where the strong
body musculature results in highly contractile
zooids. The development of the musculature,
the length of the oesophagus and the nature
of the common test, therefore, provide the
only morphological characters to determine
the species, and it is possible that some mis-
identification occasionally occurs. The record-
ed distribution of this species also suggests

that more than a single species is represented.

Pseudodistoma cercum Michaelsen

Pseudod'tstoina cereurn Michaelsen, 1924, p. 364. Kott,
1972a, p. 12 and synonymy.

New Records: Western Port (Crawfish Rock).
Cape Nelson (near Portland, vertical faces and
roof of cave, moderate surge, 5 m).
Distribution: The species is apparently com-
mon intertidally and from depths down to 70
metres off the South Island of New Zealand.
Other records are from the eastern coast of
Victoria, and off the South Australian coast
and from Dakar.

Description: The colony is very damaged and
its form is not discernible. The test is very
soft, jelly-like and transparent. Both apertures
are 6-lobed, there are 3 rows each of about
20 stigmata. Stomach folds are not apparent
externally, however internally its glandular wall
is interrupted in 4 places to give, the appear-
ance of folds. The zooids are short and the
thorax, abdomen and posterior abdomen are
of equal length.

Remarks: Zooids are characteristic. In view of
the Australian and New Zealand records sug-
gesting a circum-polar distribution in the south-
ern cold temperate region the record from
Dakar (Monniot, 1969) is surprising. Mon-
niot's specimens do, however, agree with the
present colonies and with those from New
Zealand. The species is delicate and is taken
from underneath ledges and in rocky locations
where some protection is available.

Dumus areniferus Brewin
(Figs. 4, 5, 6)
Dumus areniferus Brewin, 1952, p. 453.
New Records: Western Port Bay (Crawfish
Rock).

Distribution: New Zealand: Otago.
Description: The colonies form a thicket of
elongate branching stalks, club-shaped termin-
ally with the free end obliquely flattened. The
outer test is encrusted with a single layer of
sand particles giving rigidity to the otherwise
extremely delicate test. Each terminal lobe con-
tains only a single zooid. The maximum length

ASCIDIAN FAUNA OF WESTERN PORT

Oxycorynia pseudobaudinensis

1 contracted zooid, musculature not shown on

abdomen.
Eudistoma pyriforme
2 — contracted zooid, musculature not shown on

abdomen.
Atapozoa mirabilis
3 — zooid.
Dumus areniferus
4 — colony. 5 — zooid. 6 — larva.

Aplidium depressum
7 — zooid.
Aplidium lobatum
8 — zooid.

Aplidium triggiensis
9 — zooid.

Synoicum hypurgon
10 — zooid

Sidenioides tamaramae
1 1 — zooid. 1 2 — larva.

PATRICIA KOTT

of the stalks is 60 cm. Zooids are about 3
cm long, of which the long thread-like poster-
ior abdomen is about two-thirds of the total
length. Both apertures are 6-lobed and open
directly onto the surface of the terminal flat
surface of the stalks. There are no protective
flaps of test protecting the apertures such as
are found in Euherdmania australis. There are
about eight very fine thoracic muscles along
each side of the thorax. There are six long
stigmata crossed by parastigmatic vessels in
each of four rows. The abdomen is approxi-
mately the same length as the thorax. The
oesophagus is of moderate length, the smooth
walled stomach is elongate and there is a
posterior stomach and a duodenal region. A
mid-intestinal region occupies, with the
stomach and oesophagus, the descending limb
of the gut loop before it enters the rectum in
the pole of the loop. Testis follicles are pre-
sent in a single row in the posterior abdomen
and there is a group of ova anterior to the
testis lobes, a little distant from the posterior
end of the abdomen. The gonads occupy only
the posterior half of the posterior abdomen.

The larvae are 0-5 mm long and there may
be up to nine in the peribranchial cavity. They
have paired rows of ampullary vesicles dor-
sally along cither side of the endostyle and
along either side of the postero-ventral aspect
of the larval body. There are the usual three
papillae anteriorly and these alternate with
median ampulllae. Lateral ampullae are also
present either side of the median ampullae.
Remarks: The specimens conform exactly both
in colony and zooid form with those described
by Brewin from New Zealand. The species
resembles both Euherdmania australis and
Ritterella herdmania, both with a similar col-
ony. Externally it is distinguished from E.
australis by the absence of the flap of test
which protects the external aperture in the
latter species (see Kott, 1957, 1972b), and
from E, herdmania by the fan shape of the
terminal tip of each lobe. It is possible that
the aberrant colonies mentioned by Kott
(1957) that were taken with E % australis were
actually specimens of the present species.

Polyclinum marsupiale Kott
Polyclinum marsupiale Kott, 1963, p. 83.
New Records; Western Port (Crawfish Rock).
Distribution: S. Aust.: Victor Harbour; Tas.:
Hunter Island; Qld. : Great Barrier Reef
(Heron Island).

Description: Colonies are mushroom-shaped,
up to 2 inches in diameter across the upper
surface of the head which is supported on a
short stalk; or alternatively the colonies may
be spherical and sessile, fixed by a small
area of the base. There is a dense outer layer
of sand on the test absent sometimes from
parts of the upper surface. Internally the test
is very soft with only very occasional sand
grains included. The internal test is traversed
by canals in which the zooids are contained,
and forms only thin septa between these can-
als. Preserved colonies are therefore often col-
lapsed and flattened.

The zooids, opening around the upper half
of the colony, are very small. The branchial
aperture is terminal. The antero-dorsal atrial
aperture is on a short siphon with a circular
sphincter muscle and the opening is protected
by a pointed muscular languet from the body
wall anterior to the siphon. There are 14 to
15 rows of 10 to 12 rectangular stigmata with
the usual rounded papillae on the transverse
vessels. The stomach is smooth externally with
an inner glandular wall.

Remarks: The heads of living colonies are
apparently distinctively spherical, although in
preserved specimens the soft internal test col-
lapses and they are flattened and sometimes
appear lobed or folded.

Aplidium depressum Sluiter
(Fig. 7)

Aplidium depressum Sluiter, 1909, p. 102. Kott, 1963,
p. 95 and synonymy.

New Records: Western Port (Crawfish Rock,
Rutherford Channel)

Distribution: Previously recorded only from
Bargara (Queensland) and from Indonesia
and the Philippines. The species is common in
those locations from which it has been re-
corded. The reason for these isolated records
is not known.

ASCID1AN FAUNA OF WESTERN PORT

Description; Soft, jelly-like, flat investing col-
onies that are minute and circular fixed by a
small area of their base, or more extensive
fixed by the whole extent of the basal surface.
The species commonly invests stalks and
fronds of weed. Only very sparse sand grains
are enclosed in the semitransparent brownish
common test through which the zooids are
clearly evident. In the smaller colonies the
zooids are arranged in two or three circular
systems of about six zooids, but in the larger
colonies these expand into double row systems.
The zooids are minute with an inconspicuous
sessile atrial aperture halfway down the dorsal
surface of the thorax. There is a single short,
pointed atrial languet from the upper border
of the aperture. There are five rows of about
eight stigmata. The thorax and abdomen are
of equal length and together represent half
the length of the zooid. The stomach has 11
distinct folds.

Remarks: The small number of rows of stig-
mata, with the number of stomach folds, the
form of the colony and the nature of the test
distinguish the species.

Apldium lobatum Savigny
(Fig. 8)

Aplidittm lobatum Savigny, 1816, p. 182. Kott, 1963,

p. 97 and synonymy.
Non Psammaplidium lobatum; Herdman, 1899, p. 85

«ApUdium solidum Herdman, 1899; Millar 1963;

>A. arboratum Kott, 1963).

New Records: Western Port (Crawfish Rock).
Distribution: Florida, West Indies, the Mediter-
ranean, Red Sea, Indonesia, Queensland and
the Great Barrier Reef and New South Wales.
The present record extends the southern range
of this species from the east coast of Australia.
Description: The colonies are irregular and
investing. The common test is firm and hard
with sand throughout. Zooids are minute with
the thorax, abdomen and posterior abdomen
all of equal length. The sessile atrial aperture
has a deeply divided trifid languet from the
anterior border of the opening. There are 8
fine longitudinal muscle bands along each side
of the thorax extending onto the abdomen and
posterior abdomen. There are six rows each of

about six stigmata. The four stomach folds
are only apparent internally.
Remarks: The species appears to be adapted
for a rigorous environment, and is found in-
vesting the undersurface of rocks and in the
present case, in algal holdfasts. The firm test
and zooids with few stomach folds, are similar
to the condition found in A, solidum Herdman
in which zooids open on both sides of flat
lamellae.

Aplidium triggiensis Kott
(Fig. 9)

Aplidium triggiensis Kott, 1963, p. 104.
New Records: Western Port (Crawfish Rock).
Distribution: W. Aust.: Rottnest Island, Triggs
Island and Nornalup; Vict.: Balnarring Beach.
Description: The colonies are very soft and
investing stones, etc. Sometimes they are pro-
duced basally into projections which extend
into or around the substrate to form a very
firm adhesion. Varying quantities of sand are
present in the colonies. Posterior abdomina
cross one another in the basal test. The zooids
are minute, the thorax is 1-3 mm long and
generally shorter than the abdomen when con-
tracted; the posterior abdomen is long and
thin and up to twice the length of the rest of
the body.

There is a sessile atrial aperture about one
third of the distance down the dorsal surface
of the thorax, with a short, pointed, undivided
languet from the upper border of the opening.
There are 9 to 10 rows of about 15 stigmata.
The oesophagus is long and the stomach, pre-
sent about half way down the abdomen, is
broken up into 14 to 15 distinct folds. A single
embryo is present in the peribranchial cavity.
It is 0-6 mm long and anteriorly there is a
multiplicity of adhesive papillae in the median
line around the anterior end of the larva as
previously described for this species (Kott,
1963).

Remarks: This species also appears to be
adapted for very rigorous conditions, both by
the form of the colony and its tendency to
produce extensions to fix it firmly to the sub-
strate. The larval form is quite distinctive and
is large in relation to the size of the zooid.

PATRICIA KOTT

Consequently, a maximum of two larvae have
been reported as present in the peribronchial
cavity. In the absence of this distinctive larval
form, these species could be confused with
Aplidium multiplicatum (Sluiter) which has
been recorded from Queensland, Japan, the
Philippines, Indonesia and from Broome,
North-western Australia (see Millar, 1963). In
the latter species, however, the posterior ab-
domen is relatively short and the testis lobes
form bunches in the posterior abdomen, rather
than double rows, as in A. triggiensis.

Aplidium pliciferum (Redikorzev)

Atnaroucium pliciferum Redikorzev, 1927, p. 390.
Kott, 1972a, p. 13 and synonymy.

New Record: Western Port (Tankerton Jetty).
Distribution: See Kott, 1972a.
Description: The colony is a firm gelatinous
cushion with a flat upper surface. There is
a short stalk from the middle of the under
surface. The margin of the colony is rounded.
The test is semi-transparent and there is neither
encrusting nor included sand. The zooids are
tightly packed in double rows radiating from
common cloacal apertures randomly placed
on the upper surface. Anteriorly the zooids
are parallel to one another and vertical,
although the posterior abdomina may be more
irregularly orientated in the basal half of the
test. There are 12 fine longitudinal thoracic
muscles extending separately along the abdo-
men and both sides of the posterior abdomen.
They are never gathered into a close band.
The atrial aperture is sometimes produced into
a short siphon and the pointed single bifid lip
extends from the upper border of the aperture.
There are 11 — 15 rows each of about eight
stigmata. The thorax and abdomen are about
the same length and the posterior abdomen
is long and threadlike. The stomach has 18
to 20 regular longitudinal folds.

Larvae are present in the peribranchial cav-
ity. They have a double row of ampullary
vesicles from the lateral ridges on either side
of the median papillae which alternate with
median ampullae.

Remarks: Although there is some variation in
the number of rows of stigmata, in the number

of stomach folds and in the shape of the
colony, the firm gelatinous flat-surfaced colony
form varies only in relation to the area by
which it is fixed. The larvae are also character-
istic. The species differs from the closely re-
lated A, flavolineatum which has more thoracic
muscles, more stomach folds, lateral branches
on the larval median ampullae and no larval
ampullary vesicles.

Synoicium hypurgon (Michaelsen)

(Fig. 10)

Macroclinum hypurgon Michaelsen, 1924, p. 401.
Synovium hypurgon; Kott, 1963, p. 86 and synonymy.

New Records: Western Port (Crawfish Rock).
Distribution: W. Aust.: Rottnest Island, Fre-
mantle; Great Barrier Reef: Heron Island.
New Zealand: North Island.
Description: The present specimen consists of
three large clavate to mushroom shaped lobes
joined basally and to varying extends along
their sides to form a large, hemispherical
colony, 5 cm in diameter and sessile basally.
There is sand around the sides of each lobe
but not on the upper surface where the zooids
open. The test is soft and gelatinous and has
no foreign bodies. The zooids are present in
the outer layer of the upper surface. There is
a small, sessile atrial aperture one third of the
distance down the dorsal surface of the thorax
with a large, triangular atrial languet rising
from the body wall anterior to the aperture.
There are eight longitudinal muscle bands on
the thorax. The branchial sac is very long
and narrow with 13 rows of eight small oval
stigmata in each row. The gut loop is short,
about half the length of the thorax. The
oesophagus is especially short and the stomach
small and smooth. There is a duodenal en-
largement and a posterior stomach in the loop
of the gut.

Remarks: Although there is considerable vari-
ation in the form of colonies of this species
and some variation in the amount of sand and
other material which is contained in the com-
mon test, the small zooids, long, narrow
branchial sac and relatively short abdomen,
together with the relative position of the atrial
tongue from the body wall rather than from

ASC1DIAN FAUNA OF WESTERN PORT

the anterior border of the atrial aperture dis-
tinguish it.

Synoicium sp.?
Record: Western Port (Crawfish Rock).
Description: The specimen is damaged and
torn, although the fragments appear to re-
present a fairly thin investing colony. The
test is semi-transparent and very soft. Zooids
appear to be arranged parallel to one another
and vertical to the upper surface. Zooids are
fairly small and the thorax and posterior ab-
domen are about equal in length, while the
abdomen is shorter. There are about 10 long-
itudinal thoracic muscles. The atrial aperture
is sessile and there is a single pointed languet
from the upper margin of the opening. There
are three rows of about 10 long rectangular
stigmata, each row crossed by parastigmatic
vessels. Dorsal languets are present in the
mid-dorsal line opposite both transverse ves-
sels and parastigmatic vessels. The stomach is
shield-shaped and smooth without any areol-
ations, although it has a glandular appearance.
Remarks: The parastigmatic vessels in the
branchial sac are unusual, although they have
previously been described for Synoicium
atopogaster Kott, 1962. The small number of
rows of stigmata in the branchial sac suggests
a relationship with Synoicium bowerbanki,
which has, however, a longer oesophagus and
a distinct atrial siphon. Further, in the present
specimen, the dorsal languets opposite the
parastigmatic vessels as well as the primary
transverse vessels suggests that the rows of
stigmata are in the process of subdividing and
in fact the most posterior row does contain
a few stigmata which are bisected in the region
of the parastigmatic vessel. Synoicium papil-
liferum differs from the present specimen in
the presence of a long siphon, although it has
a short oesophagus, as well as the same
number of longitudinal muscles and about the
same number of stigmata in each row, as
does the present specimen. It is most probable,
therefore, that this represents a juvenile of
some species of Synoicium, rather than a new
species characterised by 3 rows of stigmata
crossed by parastigmatic vessels.

Sidneioides tamaramae Kesteven

(Figs. 11, 12)

Sidneioides tamaramae Kesteven, 1909, p. 277. Kott,
1957, p. 104.

New Records: Western Port (Crawfish Rock).
Distribution: N.S.W.: Tamaramae Bay.
Description: The colonies are soft and pillar-
like lobes. The free end of each lobe is raised
into a rounded marginal ridge surrounding a
terminal depressed surface from the centre of
which there is a protruberant common cloacal
aperture. The branchial apertures are made
conspicuous by the absence of sand, around
them. They open onto rounded swellings on
the marginal ridge. The external test is com-
pletely encrusted with sand, absent only from
the region around the apertures. The test is
otherwise very soft. The abdomen is about
half the size of the long thorax. The atrial lip
is narrow and fleshy but very long with about
10 fine muscles extending along its length. The
longitudinal thoracic muscles extend along the
ventral side of the abdomen and the dorsum
of the posterior abdomen causing it to curve
when the muscles are contracted. There are
17 rows of stigmata with 18 stigmata in each
row. There is no sign of papillae on the
transverse vessels. The stomach is oval with
mulberry-like glandular swellings in its wall.
There is a duodenal region, a posterior stomach
and a mid-intestine which expands into the
rectum before it curves into the ascending
limb of the gut loop. The ovary is developed
in the thoracic wall at about mid-thoracic
level and projects into the peribranchial cavity
just to the right of the mid-line, the vas
deferens and the distal part of the rectum. The
anal opening is opposite the 7th row of stig-
mata.

There are about 18 eggs at varying stages
of development in the ovary. Free eggs are
also present in the peribranchial cavity together
with up to 20 developing embryos. Mature
embryos are 6 mm long and the tail is
wound completely around the bodv. There is
a double row of vesicles along either side of
the mid-dorsal line and a cluster of vesicles
postero-ventrally on each side of the body.
Paired lateral ampullae alternate with the three

PATRICIA KOTT

anterior papillae but there arc no median
ampullae.

Remarks: This record has extended the range
of this interesting species, previously regarded
as endemic to a small region on the coast of
New South Wales.

I rididemnuin cyclops Michaclscn

(Fig. 13}

Trididannum cyclops Miehaelscn, [921, p. 19. Kott,
L966, p. 286 and synonymy. Eldredge, 1967: 183.

New Records: Western Port (Flinders Jetty,
Eagle Rock).

Distribution: West Indian Ocean. N. Aust. :
Darwin, Great Barrier Reef.
Description: Both the present records represent
extensive colonies, almost completely investing
specimens of Ascidia sydneyensis. h\ both
cases the branchial aperture is free, although
in one specimen the didemnid has grown over
the atrial siphon, leaving a small space be-
tween the test of the host through which the
excurrent water could flow.

The surface of the colony is smooth with
a superficial layer of flat bladder cells and
some spherical purple pigment cells. The spi-
cules are dense beneath the layer of bladder
cells and in the thoracic region and become
less dense toward the base of the colony where
they are absent altogether. They are from 003
— 005 mm in diameter with up to 12 pointed
rays in optical section. There are no zooxan-
thellae in the common cloacal system of these
specimens. There is a very shallow thoracic
common cloacal cavity. The zooids
have a minute thorax with three rows of
stigmata. There is no endostylar pigment cap
present in these specimens. The retractor
muscle is fairly long. There is no atrial siphon,
although there is a well defined and fairly long
anterior lip from the border of the aperture.
There is a single undivided testis follicle with
8 J coils of the vas deferens.
Remarks: The surface bladder cell layer, the
shallow thoracic common cloacal canal, the
absence of an atrial siphon and the form and
distribution of the spicules are characteristic
of this species. The extensive colonies are

different to the typical small colonies of tropi-
cal specimens. The absence of zooxanthellae
should be especially noted as these are invari-
ably present in specimens previously described.
It is possible that the zooxanthellae are associ-
ated with a tropical environment and should
not be regarded as a specific character. In
addition, the endostylar pigment cap has in-
variably been present in previously described
specimens of this species, but it is possible
that its presence is a variable character as in
7'. cerehrijorme (see below). However, these
specimens do diverge from the characteristic
facies of this tropical species and it is possible
that there is a cline in its characters that is
evident at the southern limit of its range.

Trididcmnum cerebriforme Hartmeyer

(Fig. 14)

Trididcmnum cerehrijorme Hartmeyer. 1913, p. 139.
Kott, J972d, p. 247; 1972e, p. 47 and synonymy.
non i rhthlcnuutm cebrijorme\ Kott, 1972b, p. 178.

New Records: Western Port (Crawfish Rock).
Distribution: South and West Africa; Indian
Ocean; S.W. Aust.; S. Aust.; Vict: Phillip
Island; N.S.W.; Qd.; Gulf of Carpentaria. It
therefore has a wide distribution in the south-
ern temperate to subtropical regions and is
absent only from the eastern Pacific and the
Western Atlantic.

Description: The colony is irregularly lobed.
Branchial openings are conspicuous and slightly
protruberant, owing to the density of spicules
rilling the branchial lobes. There is the usual
posteriorly directed atrial siphon. There is an
extensive posterior abdominal cloacal system
formed by canals traversing a central core of
test. The spicules are less dense than at the
surface. They are large, from 04 to 007 mm
in diameter with five conical rays in optical
section. The endostylar pigment cap is absent.
Remarks: With the exception of the endostylar
pigment cap the zooids and colony of this
specimen are identical with those previously
described. The pigment cap is also absent from
the specimens of T. cyclops from this locality.
Specimens from South Australia (Kott, 1972b)
without a posteriorly directed atrial siphon are
incorrectly assigned to this species.

ASCIDIAN FAUNA OF WESTERN PORT

?Didemnum candidum Savigny

(Figs. 15, 16)

Didemnum candidum Savigny, 1816, p. 194. Kott,
1972a, p. 19 and synonymy; 1972b, p. 179.

New Records: Western Port (Crawfish Rock).
Distribution: Cosmopolitan (see Kott, 1972a).
Discription: Flat small pinkish colonies, the
colour being due to the fairly sparse distri-
bution of spicules allowing the zooids to show
through. Spicules are mostly in the surface
and basal test. There is an extensive thoracic
common cloacal system. Zooids are about one
mm long with four rows of stigmata. The
anterior border of the atrial aperture is pro-
duced into an atrial lip, forked terminally.
There is a single testis follicle with 4i coils
of the vas deferens. Spicules range from 02
to 05 mm in diameter with conical pointed
to needle-like rays.

Remarks: The present young colonies have the
spicules typical of this species together with
the extensive thoracic common cloacal cavity.
Zooids of more typical colonies are brown.
The production of the anterior border of the
atrial aperture into a lip is another character
not usual for the species.

Didemnum moseleyi (Herdman)

(Fig. 17)

Leptoclinium moseleyi Herdman, 1886, p. 272.
Didemnum moselyeyi, Kott, 1972a, p. 19 and syno-
nymy; 1972b, p. 179; 1972d, p. 249.

New Records: Western Port (Crawfish Rock,
Eagle Rock).

Distribution: Pacific and Indo-Malayan region;
circum- Australian.

Description: Common, although not very num-
erous. White investing colonies and small
circular colonies on weed are available. Spi-
cules are dense throughout the test which is
rather brittle. The common cloacal canals
are thoracic and zooids are enveloped in an
independent thoracic sheath where the thorax
crosses the common cloacal cavity. The sur-
face layer of test is very thin indeed. In some
colonies there are primary canals extending to
abdominal level and surrounding discrete
clumps of zooids which are embedded abdom-
inally in the common basal test, although their

thoraces are separate, each in a discrete thor-
acic sheath. The surface layer of test in these
colonies is depressed over the deep primary
cloacal canals giving a cauliflower appearance
to the surface of the colony. Large common
cloacal openings are distributed randomly over
the surface and some spicule filled papillae
are also present on parts of the colony. The
spicules are 001 to 003 mm in diameter
with about seven pointed rays in optical sec-
tion. The common cloacal system in this
species does not appear to develop by proli-
feration of double row systems but by a de-
velopment of the primary cavity to envelop the
thorax of each zooid as it is added to the
system. Zooids are minute and colourless in
formalin. Small spherical lateral organs are
present either side of the thorax opposite the
most posterior row of stigmata. There is a
long retractor muscle. The vas deferens coils
9i times around the undivided testis follicle.
In one specimen large spherical vesicular
cells are present in the surface test surrounding
each branchial opening to form wide inter-
secting circles that interrupt the dense spicules
so that the surface appears to be pitted rather
than smooth. The spicules, cloacal system and
zooids otherwise conform with D. moseleyi.
Remarks: The vas deferns in these colonies
has more coils than is usual for the species
although a wide range has been reported. The
distribution and form of the rather constant
spicules and the size and position of the
lateral organs have been used to determine
this species and to distinguish it from Didem-
num candidum (see Kott, 1972a) which has
a similar cloacal system and a similarly wide
range recorded for the spirals of the vas def-
erens.

Didemnum patulum (Herdman)
(Fig. 18)

Leptoclinum patulum Herdman, 1899, p. 92.
Didemnum patulum Kott, 1972a, p. 18.

New Records: Western Port (Crawfish Rock,
on Ecklonia holdfasts and investing other as-
cidians; Eagle Rock).

Distribution: N.S.W.: Port Jackson; S. Aust:
St. Vincent Gulf.

PATRICIA KOTT

Description: Colonies form large sheets. The
surface is smooth. Smaller colonies may be an
even, greyish colour, but larger colonies always
have grey-blue to black mottled markings.
There is a surface layer of bladder cells and
beneath these numerous stellate pigment cells
are distributed amongst the spicules to form
the mottled markings that characterise the
species. Spicules gradually become less dense
toward the base of the colony. The pigment
cells are especially concentrated in the test
overlying the cloacal canal; occasionally they
may extend into the test beneath the surface
layer and beneath the common cloacal cavities
and when this occurs the colony is almost
black in colour. The spicules are stellate, with
pointed or rounded rays. The majority of
spicules are 03 - 004 mm in diameter with
about seven rays in cross section. There are,
however, less common spicules of similar form
but larger diameter, up to 005 mm. There
are also smaller spicules with up to 12 sharply
pointed rays in cross-section similar to those
found in Didemnum candidum.

The zooids are embedded in the rather solid
common test and open on both sides of
cloacal canals. Primary cloacal canals some-
times extend the whole length of the zooid
and may extend slightly posterior abdominally.
The secondary cloacal canals remain at the
level of the thoraces. The surface test is thick
and there is a long branchial opening. The
upper border of the atrial opening is some-
times produced into a lip. There are four rows
of about eight stigmata. The basal layer of test
in which the abdomina of the zooids are
embedded is rather thick and gelatinous.
Remarks: This species is the most common
ascidian in the particularly rich ascidian fauna
at Crawfish Rock. The characteristic marking
caused by greyish-black stellate pigment cells
overlying the common cloacal canals and the
thick layer of basal test distinguish the species
from Didemnum candidum in which there is
the same variety in form of the spicules. The
common cloacal system is also distinctive in
that the thoraces of zooids are not completely
enveloped bv the cloacal cavity as in D. can-
didum and D. moseleyi but remain embedded

in common test opening into the cloacal canals
from their dorsal surface. Although the species
has been recorded from Port Jackson and from
St. Vincent Gulf it is never present in the same
high density as at Crawfish Rock and it has
never been reported from Port Phillip Bay.

Didemnum turritum Michaelsen
(Figs. 19, 20)

Didemnum turritum Michaelsen, 1930, p. 521. Kott,
1962, p. 319.

New Records: Western Port (Crawfish Rock;
Eagle Rock).

Distribution: S. W. Aust.; S. Aust.: St. Vincent
Gulf.

Description: Pinkish investing colonies. Large
cloacal apertures are scattered over the surface.
The branchial apertures are also conspicuous
owing to the density of spicules in the test
covering the branchial lobes. A single lobe
of the branchial aperture sometimes developes
a hollow pointed papilla from its base. Small
rounded pigment cells line the common cloa-
cal cavities. The surface of the test may be
depressed over the deep primary common
cloacal canals to form furrows on the surface.
Clumps of zooids are surrounded by these
deep primary canals which sometimes extend
posterior to the abdomina of zooids. The
secondary cloacal cavities are thoracic. The
spicules are regularly stellate with about seven
conical rays in section and are 0-3 to 0-4
mm in diameter. The thorax of each zooid
is about 10 mm long with large oval lateral
organs which occupy a pronounced pit in the
thoracic wall along most of its length. The
branchial siphon has a well defined circular
sphincter muscle but is not very long although
the surface layer of test is thicker than that
of either D. candidum or D. moseleyi. The
atrial aperture is extensive exposing most of
the dorsal part of the branchial sac and some-
times its anterior border is produced into a
pronounced forked lip. The thoracic retractor
muscle was not detected.

There are four rows of about eight stigmata.
The gonads were not distinguishable.
Remarks: The species is readily recognized by
the hollow pointed papillae associated with

ASCIDIAN FAUNA OF WESTERN PORT

Trididemnum cyclops
13 — spicules.

Tridemnum cerebriforme
14 — spicules.
Didemnum candidum
1 5 — zooid . 1 6 — spicules.
Didemnum moseleyi
17 — spicules.
Didemnum patulum
18 — spicules.
Didemnum turritum
19 — thorax. 20 — spicules.

Didemnum augusti

2 1 — thorax. 22 — spicules.

Didemnum roberti

23 — spicules.

Didemnum spongioides

24 — spicules. 25 — larva.

Lissoclinum fragile

26 — spicules.

Diplosoma translucidum

27 — zooid. 28 — diagrammatic cross section through

colony.
Polysyncraton victoriensis
29 — spicules.

PATRICIA KOTT

one of the branchial lobes of each aperture
in certain limited areas. These papillae super-
ficially resemble those sometimes occuring in
D. moseleyi although in the latter species
they are not hollow and are not specifically
associated with the apertures. The thicker
surface test, the relatively large zooid and the
large oval lateral organs also distinguish the
species.

The hollow papillae protecting the branchial
apertures are reminiscent of those in D. nek-
ozita Tokioka, 1967, from the Palau Islands
and the Philippines. The latter species, how-
ever, has distinctive spicules and a thoracic
cloacal system.

Didemnum augusti Michaelsen

(Figs. 21, 22)

Didemnum augusti Michaelsen 1920, p. 39. Kott,

1962, p. 323; 1972d, p. 247.
1 Didemnum partiturn Tokioka, 1953, p. 191.

New Records: Western Port (Crawfish Rock).
Ram Head (18 miles south of Mallacoota
Inlet).

Distribution: S.W. Aust.; S. Aust.: Reevesby
Island; Vict: Balnarring Beach; West Indian
Ocean.

Description: Very extensive, thin, investing
colonies with dense white spicules, less dense
only in the basal test. The spicules are stellate
from 03 to 05 mm with 5 to 7 conical
pointed rays in optical cross section. The
surface of the test is furrowed and has a
cauliflower-like appearance where the surface
test is depressed over the deep primary can-
als. The primary canals extend the whole length
of the zooids between pillars of common test
in which the abdomina are embedded. Only
the dorsal aspect of the thorax is exposed to
the common cloacal canals. Some secondary
canals are present but the thorax is never
enclosed in its own discrete sheath of test. The
test along either side of the atrial opening is
thickened but there is no lateral organ. The
thorax is small, 0-6 mm, with four rows of
stigmata. There is a retractor muscle always
present.

Remarks: The species is distinguished from
D. turritum and D. moseleyi, which often have

the same surface furrows and deep primary
canals, by the solid pillars of test in which
the zooids are embedded, by the very small
thorax, the absence of a distinct lateral organ
and by the large spicules with few conical rays.

Didemnum roberti Michaelsen
(Fig. 23)

Didemnum roberti Michaelsen, 1930, p. 516.
Didemnum ternatanum; Kott, 1972b, p. 179.

New Records: Western Port (Crawfish Rock;
Eagle Rock).

Distribution: W. Aust.: Shark Bay; S. Aust:
Elliston Bay.

Description: Investing colonies with a smooth
surface and dense spicules in the surface and
basal layers of test. The common cloacal aper-
tures have their borders stiffened with spicules
and are very conspicuous. Some colonies are
flattened but in some, rounded lobes are
developed by a thickening of the basal test
to form a central core of test. There are ex-
tensive posterior-abdominal cloacal spaces and
the zooids are suspended in clumps between
the surface and basal layers of test, anchored
basally by a short single strand of test, and
at the surface by the branchial lobes of re-
spective zooids. Secondary common cloacal
cavities surround the thoraces of the zooid,
each surrounded by a discrete layer of test
and with a large lateral organ occupying most
of each side of the thorax (as in D. turritum).
The spicules are not so thick in the test
surrounding the zooids. The surface test is
fairly thick, again resembling D. turritum.
Spicules are almost spherical, 002 to 004
mm in diameter with rounded rays. The zooids
are small. The testis follicle is undivided and
has 7i coils of the vas deferens around it.
Remarks: The present colonies diverge from
Michaelsen's (1930) specimens only in the
presence of spicules throughout basal or axial
test. In the Shark Bay material the basal
layer of spicules was confined to a layer
beneath the posterior abdominal canals to
form a sort of endoskeleton.

Colonies from Elliston Bay (Kott, 1972b)
which are identical with the present colonies
from Western Port, were erroneously assigned

ASCIDIAN FAUNA OF WESTERN PORT

to the species Didemnum ternatanum Gotts-
chaldt. Although the three-dimensional com-
mon cloaca and the size and form of the
spicules are similar to those of D. ternatanum,
the present species is distinguished from it
by its external, oval lateral organ, by the more
densely distributed spicules; by the multiplicity
of common cloacal apertures and extensive
colony; and by its firmer consistency. D. rob-
erti is distinguished from D. bistratum Michael-
sen, 1920 from West Africa by the form of
its spicules (those of the latter species are
spherical and hollow) and by its external
lateral organ. D. spongioides also has a simil-
arly labyrinthine common cloaca, but its spi-
cules are stellate, with fewer, conical rays, and
fewer coils of the vas deferens.

Didemnum roberti has previously been
described as yellow, or yellowish— no inform-
ation is available on the in vivo colour of
the present colonies.

Didemnum spongioides Sluiter

(Figs. 24, 25)

Didemnum spongioides Sluiter, 1909, p. 67. Kott,
1962, p. 318; Eldredge, 1967, p. 193.

New Records: Western Port (Crawfish Rock).

Distribution: Caroline Is.; Indonesia; W. Aust.:

Rottnest Island; Tas.: Oyster Bay. The records

suggest a circum-Australian distribution.

Description: Colonies are rounded to conical

with a terminal common cloacal cavity. The

test is firm. Spicules are present in a layer

beneath a surface layer of bladder cells at the

level of the branchial siphons. They are less

dense beneath this layer and are entirely

absent from the test core that occupies the

centre of the colony. The surface of the test

is covered with minute spicule-filled pointed

papillae that project through the bladder cell

layer and, when magnified, give to the surface

a spotted appearance.

The spicules are stellate with about seven

conical rays in cross section, and range from

002 to 006 mm in diameter. An extensive

common cloacal cavity separates the outer

spicule and zooid bearing layer of test from

the inner spicule free test core in which

embryos develop. Cloacal canals extend into

the zooid bearing layer but these do not
separate clumps of zooids from one another.
The openings of the common cloacal canals
into the posterior abdominal chamber are
shown by Sluiter (1909, Plate 6, fig. 9) and
the ridges and trabeculae he describes are
formed by the roof of the cloacal chamber
enclosing abdomina of zooids projecting into
the chamber, between the openings of the
canals. These ridges and trabecula are not
the imprint of the substrate on the base of
the colony as Eldredge (1967) has suggested.
Zooids are small. The thorax, when contracted
is only 0-5 mm long. There is a wide atrial
opening and a small rounded lateral organ
opposite the 4th row of stigmata. The vas
deferens coils 6i times around the undivided
testis follicle.

Embryos are packed in the central test core
at the base of the common cloacal chamber,
into which they move through the occasional
strands of test that connect the surface layer
to the central core. They are 0-9 mm long
when mature, have an ocellus and an otolith,
and four pairs of lateral ampullae. The tail
winds once around the embryo.
Remarks: The species is related to D. lambitum
in the form of cloacal system and the spicules
and is distinguished from that species by the
presence of a bladder cell layer and by the
larva in which the ampullae are not sub-
divided.

D. spongioides; Eldredge, 1967, differs from
the present specimens in the presence of pig-
ment cells, the even investing form of the
colony and in the condition of the cloacal
system with well developed thoracic secondary
canals and primary canals extending postero-
abdominally but not forming a continuous
space separating surface from central or basal
test. The thickness of the surface layer of
test, the arrangement of cloacal canals, the
spicule form, size and arrangement, and the
presence of pigment cells of Eldredge's speci-
mens are identical with those of D. turritum
from which they differ only by the absence
of hollow pointed papillae on the surface.

The colony is typically "sponge-like" in
external appearance, rounded and sessile.

PATRICIA KOTT

Didemnum lambitum (Sluiter)

Didemnoides lambitum Sluiter, 1900, p. 18.
Didemnum lambitum; Kott, 1962, p. 317 and syno-
nymy; 1971, p. 19; 1972a, p. 18.

New Record: Port Phillip Bay (Hobson's Bay).
Distribution; N. Z.: Chatham Island, North
Island, South Island, Stewart Island; Tas.; S.
Aust. ; St. Vincent Gulf.
Description: The colonies are more or less
fan shaped, made up of vertical lamellae or
columns. These may fuse for the greater part
of their length, or only basally or terminally.
The free outer edge of the fan is more or less
flattened. Common cloacal apertures are large
and rounded and are occasionally but not
always found on the free ends of the lobes.
The test is firm and gelatinous, without sand.
There is a central gelatinous core of test
surrounded by specially extensive common
cloacal spaces. The zooids are small and num-
erous, closely packed in the outer layer of test.
There are 8i coils of the vas deferens around
a single undivided testis follicle.
Remarks: This species appears to be limited
to the more temperate waters of Australia
and New Zealand extending north only to
N .S. W. on the east coast of Australia. The
spicules are usually present in the surface test
at the level of the zooids but are often absent
in the remainder of the test. In one of the
present colonies spicules are absent entirely.
The relationship of the present species to Z>.
spongioides is close. Both have a firm gel-
atinous test and a similar common cloacal
system. In both the spicules are usually absent
from the central test core and form a layer
only at the level of the branchial siphons. In
both they are stellate with about seven conical
pointed rays. The species appear to be distin-
guished only by the absence of a superficial
bladder cell layer in D. lambitum and by the
larvae which, in the latter species, has sub-
divided lateral ampullae. Generally the col-
onies of D. lambitum are higher than those
of D. spongioides. D. spongioides has been
recorded from the tropics but D. lambitum
has not. Tt is possible that both species repre-
sent different stages in development of a single
species, however, additional specimens, to-

gether with larvae will be needed to resolve the
question.

Didemnum skeati (Sollas)

Hypurgon skeati Sollas, 1903, p. 729.

Didemnum psammatodes var. skeati; Michaelsen, 1920,

pp. 22, 27 and synonymy. Hastings, 1931, p. 95,

Kott, 1962, p. 326.

New Records: Western Port (Crawfish Rock;
Eagle Rock).

Distribution: Malaysia; Indian Ocean; Vict.:
Flinders; Qd.: Moreton Bay, Sarina, Low Isles;
Torres Strait: Possession Island. The species
has not been recorded from Western Australia
but in view of its Indian Ocean occurrence
could be expected to occur there.
Description: A large number of extensive
sheets, blackish in colour owing to embedded
balls of mud throughout the test. Small groups
of spicules, as previously described, are pre-
sent over each branchial aperture. The cloacal
canals are thoracic. Zooids are very small.
Remarks: The specimens conform completely
with previous descriptions. In view of the
constant nature of this form and its consistent
differences from D. psammatodes, it has been
elevated to specific rank.

Lissoclinum fragile (Van Name)
(Fig. 26)

Diplosomoides fragile Van Name. 1902, p. 570.

Lissoclinum fragile', Eldredge, 1967, p. 245 and syno-
nymy.

? Diplosoma (sic) caulleryi Ritter and Forsyth, 1917,
p. 489.

? Lissoclinum caulleyri; Van Name, 1945, p. 114.

? Lissoclinum marpum Millar, 1953. p. 301.

? Lissoclinum bilobatum Millar. 1955, p. 180.

? Lissoclinum japonicum Tokioka, 1958, p. 73.

? Lissoclinum notti Brewin, 1958, p. 457.

Neyv Records: Western Port (Eagle Rock)
Distribution: West Indies; ? East Africa; ?
South Africa; ? Japan; Pacific; ? California;
? New Zealand. Except for Lissoclinum fragile
Van Name and Lissoclinum caulleryi Ritter
and Forsyth, the suggested synonyms are known
only from single records. The present speci-
men is the only record from Australia. The
lack of records may be explained bv the brittle
nature of the very thin investing colony which
is removed from the substrate only with the
greatest difficulty. The species is probably

ASCIDIAN FAUNA OF WESTERN PORT

circum-tropical and extends into temperate
regions of both the northern and southern
hemispheres.

Description: The colony is thin and extensive,
investing a very large specimen of Ascidia
sydneysis. There are pinkish brown pigment
cells in the surface test. Spicules are very
dense throughout, and the colony is very
brittle. Spicules do not line the branchial lobes
and are absent from a circular area in the
region of the branchial aperture through which
the interior of the colony is visible. They are
small, 02 to 03 mm in diameter, burr-like
with many flat ended rays. The cloacal cavity
is mainly thoracic, primary canals sometimes
extend to the abdominal level but never pos-
terior to the zooids. The abdominal portion
of the zooids is embedded in the basal test
which is very solid and hard owing to the
density of spicules. The thoraces cross the
cloacal cavity in independent test sheaths that
are interrupted over the dorsal surface and
most of each side of the thoraces they en-
velope, A very large 'flap like' lateral organ
is present supported on the edge of the test
sheath near the ventral border of the zooid
and overlapping the branchial sac opposite the
interval between the third and fourth rows of
stigmata. There are four rows of stigmata with
eight stigmata in each row. There are two
testis follicles with a straight vas deferens,
hooked proximally around between the two
testis follicles.

Remarks: The synonymy suggested above was
first indicated by Kott (1962). The specimens
all have a deeply indented atrial aperture, a
similar two dimensional cloacal system, and
similar spicules within the same size range
although there is some variation in their den-
sity and arrangement. The lateral organ is
usually present and is flap-like and opposite
the third to fourth rows of stigmata in all cases
except in L. fragile; Tokioka, where it appears
to be elliptical, not supported on the edge of
the test sheath and opposite the second row
of stigmata. In L. fragile; Eldredge, it is des-
cribed as a "small flap-like" organ and in
L. marpum Millar it is also small. In L. fragile
Van Name it is not always present and hns

not been described at all for L. notti Brewin.

Its presence and degree of development is,
therefore, apparently variable and its use as
a distinguishing character would not in any
case resolve the taxonomy of the forms in-
dicated above on any rational geographic
ground.

Larvae have been described for L. notti
Brewin and L. fragile; Eldredge. They are
identical in size and form although some of
Eldredge's specimens had a layer of small
opaque particles surrounding the larval body
similar to the particles described for L. os-
irearium; Kott (1962).

The present species differs from L. ostrear-
ium Michaelsen and L. molle Herdman in the
absence of a three dimensional cloacal system.

Lissoclimim osfrearium Michaelsen

Lissoclinum osfrearium Michaelsen, 1930, p. 526.
Kott, 1962, p. 308 and synonymy.

New Records; Western Port (Crawfish Rock;
Flinders Jetty).

Distribution: W. Aust.: Rottnest Island; S.
Aust.: St. Vincent Gulf; Qd.: Great Barrier
Reef.

Description: The colonies are very thin invest-
ing and rather delicate. There are some black
pigment particles in some parts of the colony.
There is a thin layer of surface test. The basal
test is thicker, sometimes enclosing abdomina
but more often clumps of zooids are anchored
to the basal test by a single narrow strand of
test. The common test then subdivides to
enclose each zooid in an independent test
sheath for almost its whole extent and anchor-
ing it anteriorly to the surface test. Each in-
dividual test sheath is interrupted dorsally to
expose a large part of the dorsal surface and
sides of the branchial sac to the extensive
common cloacal cavity. The spicules are dis-
tributed in varying density in different parts
of the colony and are often almost entirely
absent. They are less dense in the surface
layer than in the remainder of the test. The
spicules are 0025 to 003 mm in diameter
and are characteristic, with a large number
of flat-ended rays. There is a small, flap-like
lateral organ opposite the interval between

I \ I'A'I'KICIA KOTT

the third and fourth rows of stigmata. No developed and zooida tend to remain in

embryos are present in thc&e Victorian colonies, clumps. This also gives the colony a firmer

Remarks: The atrial aperture and the lateral consistency.

organ are similar to ihose of L. fragile. They

arc distinguished only by the colony which ,^. , . *„ , A • i

' ■ , r , Dinlosoiiiii niyncn MaeDonald

demonstrate!! a maximum development oi the

cloacal system as in other species of Ltssoc- Diplosoma rayneri MaeDonald, IKV>, p. $73.

/ i / »■ / Ltptocllnurn i Leptocllnuni ) rayneri: Koii, 1966, p.

Imum and Uiplosoma. \ tH) v * '

Eldrcdgc ( 1967) drew attention ^o the fact Diplosoma Hstgranum; Rowd, 1966, p. 458 and syno-
nymy.
Diplosoma macdonaldt\ Eldredge, 1967, p. 231.

that the difTcrence in the shape of the stigmata

is probably nol a valid character to distinguish

/ fragile and L, ostreartum, it is possible New Records; Western Port (Crawfish Rock).

that the shape of the stigmata is affected by Distribution: Cosmopolitan (see Rowc, L966).

the extent to which colonies with dense Spi- Description: Typical delicate colonics. Veg-

cnles retain their original shape when pre etativc reproduction in progress. No mature

served in formalin so thai the branchial site gonads observed.

is maintained in an extended condition by the

rigid test „ . . ■ • ■ i tl

Polysyncraron ormculum Kott

Polysyncraton orblculum Kott, 1962. p. 301. Kott,

i)i|>ioM>m;i transluctdum (Hartmcyer) 1872a, p. 21,

(Figs, 27, 28) New Records: Western 1*011 (Crawfish Rock).

Uptocllnum transiucidum Hartmeyer, 1909. p. 1490 Distribution: W. Aust.: Rotlnesl Island; S.

Diplosoma transiucidum', Kott, 1962, p. 306 and Ausl.: St. Vincent (mil.

synonymy, Description: The colonies are small and in-

New Records: Western Port (Eagle Rock). vesting with the usual circle o\' large vesicular

Distribution: W. Aust.: Oyster Harbour, Al- eells around each branchial aperture. There is

bany; N. W. Aust.; Indonesia. B Single layer ot spicules in the surface test.

Description: The colony is lour,, narrow and interrupted by these large vesicular cells. The

irrcgulai investing a worm tube. It has a zooids, with the red brown pigment In them,

transparent test that is fairly soft but tough give to the colony a pinkish brown colour.

and not jelly like. The basal test is extended Sonic-times the vesicular cells are so large

upwards in a lamella along, the mid longUud- that they are almosl eonlluent. Because these

iual axis Of the colony. Strands of test from cells are transparent and interrupt the distii-

both sides o\' Ihis lamella support clumps o[ bulion o[' Spicules the surface of the test

ZOoids, The common test then subdivides to appears to be pitted, or, when they arc almost

form tesi sheaths supporting the thorax of confluent, it appears lo be depressed into a

each zooid independently at the surface. The narrow trough or furrow around each opening

zooids are fairly large with the thorax about 2 so that the apertures are at the apices o\'

nun long,. There are four rows o\~ about It) apparent mounds over each zooid. The spicules

Stigmata. A large pari oi the branchial sac ftW 02 to 003 mm in diameter and are

is exposed through the wide atrial opening, regularly stellate with about eight conical rays

There is a long rectum. Oesophageal buds in optical section.

are present, but gonads were not distinguished. The cloacal canal is shallow and thoracic,

Remarks: The species is distinguished from D. the zooids small and completely embedded.

rayneri by its firmer lest and, although the Gonads were not detected.

cloacal system is typical o\' the genus with Remarks: The form o( the colony and arrange

long test strands anchoring the /ooids basallw meat of spicules has been used to determine

the secondary cloacal spaces are not so well this species.

ASC1DIAN FAUNA OF WESTERN PORT

Polsyncraton victoriensis n. sp.

(Fig. 29)

Type location: Western Port (Crawfish Rock,
8 m, on Ecklonia holdfasts) Holotype, Nat-
ional Museum of Victoria No. H. 171.
Description: The colony forms a thin invest-
ment over weed. It is a rather dirty whitish
colour in formalin. There is a layer of bladder
cells superficially over the top of each zooid.
Between the zooids, however, spicules invade
the superficial layer of test which stands out
as spicule filled ridges between the zooids and
gives an irregular and rather angular appear-
ance to the colony. Zooids thus appear to
open into the base of furrows on the surface.
There is a very shallow common cloacal
cavity. Spicules are stellate, 003 to 006 mm
in diameter with only five conical rays in
section. They are arranged evenly throughout
the test. Zooids are small. There are four rows
each of six stigmata in the branchial sac. There
are 4] coils of the vas deferens around three
to four testis follicles.

Remarks: The arrangement of spicules is the
same as that described by Hastings, 1931 for
P. magnetae. Hastings species, however, has
smaller spicules, fewer turns of the vas defer-
ens, more testis follicles and more stigmata.

Plialhisia dcprcssiuscula (Heller)

Ascidia dcprcssiuscula Heller, 1878, p. 5.
Piiallusia; dcprcssiuscula; Kott, 1972a, p. 23 and
synonymy.

New Records: Western Port (Flinders Jetty;
Tankerton Jetty); Port Phillip Bay (Hobson's
Bay; artificial reef). Portland Harbour 6-12
metres, on rocks forming jetty.
Distribution: Ceylon; Indonesia; Arafura Sea,
Philippines; circum-Australia.
Description: The present specimens fall within
the range previously indicated for this species.
Large specimens from Portland Harbour are
black; while the smaller specimens and speci-
from Flinders jetty, up to 20 cm in length and
fixed by the whole of the left side, are brown-
ish and translucent. One of these large speci-
mens is completely invested with Lissoclinitm
fragile.

Ascidia sydneyensis Stimpson

(Fig. 30)

Ascidia sydneyensis Stimpson, 1855 (? pari), p. 387.
Kott, 1972a, p. 24 and synonymy; 1972, p. 182:
1972c, p. 237; 1972e, p. 49.

New Records: Western Port (Tankerton Jetty,
Flinders Jetty, Crawfish Rock, Eagle Rock);
Port Phillip Bay (Williamstown, Hobson's
Bay; artificial reef).

Distribution: West Jndies; South and east
Africa; Indian Ocean; Indonesia; circum-Aus-
tralian. The species is apparently circum-polar
in tropical and temperate waters of the south-
ern hemisphere although it extends north of
the tropics only to Japan.
Description: Large specimens at least 10 cm
long and up to 20 cm are available from all
stations, fixed by the whole of the left side.
The apertures are on the usual short cylindri-
cal siphons, the branchial aperture is always
turned to the left toward the substrate. The
atrial aperture from half way down the body
is turned to the right or directed anteriorly
along the dorsal surface. Sometimes the whole
antero-dorsal part of the body is turned over
to the left. The test of these large specimens
is firm and gelatinous and slightly leathery
superficially except along the left side where
it is fixed to the substrate and there it is very
thin. Two of the four large specimens from
Flinders jetty are completely covered by in-
vesting didemnids. On one specimen there is
a colony of Didemnum posteriorly and a col-
ony of Tridemnum cyclops anteriorly, which
leaves only the branchial aperture free. These
colonies overlap one another across the upper
surface in a line with the atrial aperture which
is also left free. On the other specimen the
upper surface is completely invested with a
colony of Tridemnum cyclops. There the
branchial aperture is free but the atrial aperture
is covered and the excurrent stream from the
Ascidia is apparently directed along a groove
in its test to the left of the line between the
atrial and branchial siphons and underneath
the encrusting didemnid. The gut, in all these
specimens, is filled with mud and the branchial
sac is occluded by the distended gut.

PATRICIA KOTT

Ascidia gemmata Sluiter

Ascidia gemmata Sluiter, 1895 r p. 177. Kott, 1972a,

p. 26 and synonymy.
Ascidia thompsoni; Kott, 1975, p. 10.

New Records: Port Phillip Bay (Mornington
Pier).

Distribution: Pacific; Malaysia; Indonesia;
Arafura Sea; circum- Australia. The species
thus appears to have a wide range in the Indo-
Pacific area.

Description: Two specimens are available,
about 20 cm long. The branchial aperture is
terminal on a short siphon. The atrial aperture
is sessile, two-thirds of the way along the
dorsal surface. There is a long furrow extend-
ing along the right side of the body and the
atrial aperture is directed into that furrow.
The animal is fixed by the whole of the left
side. The test is firm, gelatinous and smooth
on the surface.

Internally the siphons are more conspicuous.
There are 60 branchial tentacles, the dorsal
tubercle fills the peritubercular area. The pre-
branchial region is minutely papillated. The
dorsal lamina has the usual strong ribs on
both sides, each rib terminating in a minute
pointed tongue to form a marginal fringe. The
dorsal gland and ganglion are one third of the
distance down the body. There are intermediate
papillae in some parts of the branchial sac
in addition to the papillae at the junction of
the transverse and parastigmatic vessels. The
oesophageal opening is just posterior to the
base of the atrial opening and branchial sac
extends posteriorly to it.

Remarks: The species has not yet been record-
ed from Western Port Bay but will, very
likely, be found to occur there.

BotrvIIoides leachi (Savigny)

Botryllus leachii Savigny, 1810, p. 7.
Botrxlloides leachi; Kott, 1972a, p. 29 and svnonvmv;
1972b, p. 185; 1972d. p. 253.

.Yen- Records: Western Port (Crawfish Rock);

Portland Harbour (6 metres, on rocks from-

ing jetty).

Distribution: The species is recorded from the

north Atlantic the Mediterranean and Red

Sea, South Africa, and Indo-Australia to New

Zealand. The species does not extend into the

south Atlantic. In view of its south African
occurrence could be expected to occur more
widely in the Indian Ocean than its present
records suggest.
Description: As previously described.

BotrvIIoides nigrum Herdman

Botrxlloides nigrum Herdman. 1886, p. 50. I
1972c, p. 238 and synonymy; 1972d, p. 252.

New Records: Western Port (Crawfish Rock;
Eagle Rock). Port Phillip Bay (artificial reef;
Mordialloc). Cape Nelson (near Portland; 5
metres).

Distribution: West Indies; Red Sea; Ceylon
(?); South and East Africa; S. W. Aust.;
Vict.; N. S. \V.; Qld. The species is not as
commonly recorded as other Botrylloides spp.
Remarks: The double row systems, widely
spaced in irregular lamellae are distinctive and
it seems unlikely that the species could be
misidentified. Distribution is not cireum-polar,
as it has not been recorded from the West
African coast, nor from the Pacific Ocean.

It may be that there are two species repre-
sented, one from the Atlantic and one from
Indo-Australian waters.

Symplegma viride Herdman

Symplegma viride Herdman, 1S86. p. 144. Michaelsen,
1918, p. 101 and synonymy. Kott, 1952. r. 252 and

further synonymy; 1964. p. 129.

New Records: Western Port (Crawfish Rock.
growing on basal surface of fronds of Didem-
num patulum, encrusting a lump of tar).
Distribution: West Indies, Red- Sea. West In-
dian Ocean, Ceylon. Malaysia, the Phiiiptaes
and circum- Australian from Shark Bay (\Y.
Aust.) and south across the southern coast
to Thursday Island (off N. E. Australia).
Remarks: Colonies of this species appear to
compete with didemnids for available space
and occasionally overgrow the borders of a
didemnid colony.

Amphicarpa diptycha (Hartmeyer)

(Figs. 31-34)

Distomus diptychos Hanmeyer, l^t^. p. 87.
Ampicarpa diptycha: 19*"2e. p. 50 and further syno-
nymy.
Stolonica australis Michaelsen. 1°2", p. 202, Kott,

1972a. p. 28 and further svnonvmv; 1972b, p. 1S ; ;
1972c, p. 252.

ASCIDIAN FAUNA OF WESTERN PORT

New Records: Western Port (Crawfish Rock).
SSW. of Cape Grant (220 to 275 metres, on
a large stone); South-east of Portland (166
to 220 metres).

Distribution: North to south-western Australia;
S. Aust.; Vict.; Tas. (d'Entrecasteaux Chan-
nel). The species is known from 12 to 24
metres depth.

Description: Upright oval to elongate individ-
uals are joined to a basal membrane or on a
short stalk from basal stolons investing sponges
or stones. In some colonies individuals are
tightly packed to give a cauliflower-like appear-
ance. The test of adjacent zooids is not con-
fluent, however, and the zooids in the colony
are connected with one another only by the
basal stolon or membrane and by the adherence
of adjacent zooids to the same sand particle.
The test is covered with sand. Posteriorly where
it joins onto the basal membrane, the test may
be flattened to form a wide pseudo-stalk.
Zooids are up to 8 mm tall and 3 mm in
diameter. The branchial apertures are sessile
and a short distance down the dorsal surface.
The musculature of the body wall is strong
especially on and around the siphons but pos-
teriorly becomes weaker. The branchial folds
are high and overlapping. There are 19 rows
of stigmata. The gut forms the usual short
loop across the posterior end of the body and
the rectum continues anteriorly at right angles
to the loop. The stomach is pyriform, ex-
panded towards the pyloric end. There are
18 longitudinal folds in the stomach wall and
on the lateral aspect these folds terminate
against the sutureline and appear to be oblique
rather than longitudinal. The gastric caecum
is continuous with the suture. It extends to-
wards the pole of the gut loop and curves
around just distal to the pyloric end of the
stomach. It is tightly held against the stomach
by a body wall membrane. It is of variable
length sometimes curved or hooked or forming
one complete spiral. There is the usual liga-
ment extending between the intestine and
pyloric end of the stomach enclosing a flat
topped endocarp in the pole of the gut loop.
There is another similar endocarp between the

oesophagus and the end of the intestine where
the rectum curves anteriorly.

The anus terminates in two rounded lips.
Bisexual gonads are present consisting of a
large ovum and a wide short oviduct with a
small male follicle beneath the ovum and the
body wall, the vas deferens curving around
to open on top of the oviduct. These gonads
extend in a single row along the middle of
the right side of the body and occasionally
further towards the ventral margin. On the
left they usually extend in a line obliquely
across the anterior border of the gut loop and
along the ventral border of the body. Ovaries
only occur together with testis follicles, al-
though occasionally the male gland appears
to be spent or not to be mature. Numerous
testis follicles, however, are often scattered
in clumps antero-ventrally and postero-dor-
sally at either end of the right row of the
bisexual gonads and on the left of the body
they extend around the ventral margin anterior
to the stomach and ventral to the bisexual
gonads. This condition appears to occur in
fully developed mature zooids. In less well
developed zooids the unisexual testis follicles
are not present and the testes are confined to
the bisexual organs as described. Larvae are
present in colonies from Crawfish Rock. They
have the usual photolith and a circle of 16
ampullae anteriorly from the centre of which
three simple papillae diverge. The larval test
has a foamy vesicular appearance.
Remarks: A large number of zooids have
been examined in order to resolve the con-
fusion between this species and Stolonica
australis caused by the variable length of the
gastric caecum, variation in development and
position of the gonads and variation in the
development of the colony. The essential
differences between Stolonica australis and
Amphicarpa diptycha (viz. the density of the
zooids in the colony and the position of gonads
on the body wall) are both extremely variable
and are related to the condition and age of
the colony and the maturity of the gonads,
and the zooids. Similarly the extent to which
the basal stolons associated with S. australis
have fused to form a basal membrane and the

PATRICIA KOTT

stalk of the zooid has expanded to become
confluent with that of adjacent zooids prob-
ably depends on the maturity of the colony.
It is apparent from the zooids in the present
collection, all from the same location, that
no essential difference separates these species.
In the present specimens there is an encap-
sulated parasitic copepod in the peribranchial
cavity of most individuals.

Polyandrocarpa lapidosa (Herdman)

Goodsiria lapidosa Herdman, 1899, p. 99.
Polyandrocarpa lapidosa', Kott, 195 2, p. 250; 1972b,
p. 184; Millar, 1963, p. 730.

New Records; Port Phillip (Survey area 5;

Popes Eye, Port Phillip Heads).

Distribution: Port Phillips Heads, Western Port.

New South Wales (Port Jackson).

Description: Colony forms a large flattened

lobe with zooids opening around both sides.

The zooids conform to previous descriptions

of this species.

Polycarpa thelypanes (Sluiter)

Srylea thelypanes Sluiter, 1904, p. 68.
Polycarpa thelypanes; Kott, 1952, p. 238.

New Records: Western Port (Flinders Jetty).
South to south-east of Portland Harbour, 166
to 220 metres.

Distribution: Philippines; W. Aust.: Albany.
Kott, (1952) suggests that the species might
have been introduced to Australia with Jap-
anese oysters (Ostrea gigas) with which it was
taken. The present records, however, suggest
that this sandy inconspicuous species may
have a wider range than previously recognised.
Description: One small specimen is available
from Flinders Jetty growing on the test of a
specimen of Ascidia sydneyensis. A single
small specimen and four large specimens are
available from off Portland Harbour. The test
is very stiff and impregnated with sand. The
body wall is not very muscular and closely
adherent to the inner surface of the thin, brittle
test. The specimens are either dorso-ventrally
flattened or laterally flattened and lie on the
substrate on their right side. They are not,
apparently, fixed to the substrate but lie freely
on it. The branchial aperture rises from the
anterior end of the body and is directed at

right angles to its long axis. The atrial aper-
ture rises from about half way along the
dorsal surface. The test is tough leathery and
whitish. It is impregnated with sand every-
where except on the short cylindrical siphons.
The siphons are longitudinally furrowed and
entirely free of sand. In laterally flattened
specimens the branchial aperture is turned
over to the right towards the substrate. There
is a simple U-shaped dorsal tubercular
opening in a large peritubercular area. The
body wall is closely adherent to the test and
has very delicate though strong muscles that
radiate from the siphons but fade out on the
body. The branchial sac has four very low
rounded folds on each side of the body with
wide spaces between them. There are about
15 internal longitudinal vessels on the folds
and 9-12 between the folds. There is a very
short, rounded stomach and the gut forms a
wide open arc opening by an eight lobed anus
into the base of the atrial aperture. The gonads
are intermediate between long styelid-type
gonads and shorter polycarps. There are about
eight on both sides of the body more or less
in a row near the endostyle with others irreg-
ularly scattered over the body wall. Sometimes
the gonads are slightly curved and bent.
Remarks: The form and arrangement of gon-
ads, the course of the gut and the rigid, thin
and stiff test characterise this species. It is of
interest that the siphons remain the only con-
tractile part of this animal, the remainder of
the body wall is closely adherent to the test.
The plane along which the animal is flattened
also appears to be variable and a consequence
of its position on the substrate. The flattening
of the body contributes to the orientation of
the branchial aperture toward the substrate
and the atrial aperture away from the sub-
strate.

Cnemidocarpa etheridgii (Herdman)

(Fig. 35)

Stycla etheridgii Herdman, 1899, p. 38.
Cnemidocarpa etheridgii; Kott, 1972a., p. 31 and
synonymy; 1972d, p. 253.

New Records: Port Phillip Bay (Mornington;

ASCIDIAN FAUNA OF WESTERN PORT

Port Phillip Heads). South to south-west of
Cape Grant, (221 to 275 metres). 280° from
Cape Nelson, (220 to 294 metres); Portland
Harbour South to south-east of Portland Har-
bour (166 to 278 metres).
Distribution: W. Aust.: Triggs Island (north
of Fremantle); around the south coast of
Australia; Tag.; d'Entreeasteaux Channel;
N. S. W.: Port Jackson, Port Stephens; Qd.:
Moreton Bay.

Description: The test is whitish and tough and
leathery, but thin and paperlike in appearance.
The branchial aperture is terminal and the
atrial aperture one third to half way down the
dorsal surface. Both apertures are sessile. The
body is more or less conical in outline from
the wide rounded basal or posterior portion
and narrowing to the terminal branchial aper-
ture terminally. Posteriorly, the test may be
produced into numerous tough root-like pro-
jections or there may be a single stalk. The test
around the apertures is usually longitudinally
furrowed or sometimes, when they are with-
drawn, is transversely wrinkled. The body
wall is closely adherent to the test. The dorsal
tubercle is large and protruberant and almost
fills the peritubercular area. There is a fairly
wide, smooth margined dorsal lamina. The
branchial sac has 4 high, almost overlapping
folds on each side of the body. There is a
single large gonad in the centre of the body
wall on both sides of the body, sometimes
enclosed in endocarp-like thickening of the
body wall. The gut forms the usual long,
narrow, curved loop and also encloses an
endocarp which may cross from the left to
the right side of the body behind the branchial
sac.

Remarks: This species demonstrates a wide
range in size up to 11 cm in length. It is
characteristically large and rounded posteriorly,
with a typical tough whitish thin test. Intern-
ally the high branchial folds, the narrow
curved gut loop and the embedded gonads are
distinctive. As with Ascidia gemmata this
species has surprisingly not yet been recorded
from Western Port, but is likely to occur there.

Pyura australis (Quoy and Gaimard)

australis Quoy and Gaimard

(Fig. 36)

Ascidia australis Quoy and Gaimard, 1834, p. 614
Pyura australis, Kott, 1972b, p. 186 and further

synonymy.

New Records: Western Port (Crawfish Rock,
some fixed to a scallop shell and some with
the stalk encrusted with Amphicarpa diptycha
or attached to Ecklonia holdfasts). Port Phillip
Bay (Portsea).

Distribution: N. W. Aust.: Geraldton; around
the southern coast of Australia; Tas.: d'En-
trecasteaux Channel; S. Aust.: St. Vincent
Gulf; Vict.: Flinders.

Description: The present specimens are typical
although the projections from the test are
sometimes longer and more pointed than is
usually the case. The branchial aperture is
turned downwards toward the substrate and
protected by large tubercular extensions of the
test. There are the usual stellate siliceous
spicules 03 mm in diameter in the test
and pointed spines, less than 0-1 mm in
length, lining the siphons. The anal border
has the usual long, finger-like lobes.

Pyura cataphracta (Herdman)

(Figs. 37, 38, 39)

Cynthia cataphracta Herdman, 1899, p, 31.

Pyura multiradicata; Kott, 1952, p. 269 (part: not

Herdman's type specimen of Cynthia multiradicata

< Pyura spinifcra).

New Records: Western Port (Crawfish Rock).
South to south-west of Cape Grant, 220 to
275 metres. South to south-east of Portland
Harbour, 160 to 220 metres.
Distribution: Previously recorded only from
Port Jackson.

Description: The species is upright, slightly
narrowed toward the base. The branchial aper-
ture and atrial aperture are each on a short
siphon at opposite extremities of the upper
surface. Basally the diameter of the individual
is reduced and root-like extensions from the
test or a single stalk basally rooted may be
produced posteriorly. Externally the test is
grooved and furrowed to varying extents and
especially anteriorly. The test around the
apertures is produced into pointed processes

PATRICIA KOTT

directed anteriorly around the aperture and
depending on the extent to which the siphon
is contracted, sometimes extending across the
closed aperture. On other parts of the body,
the test may be produced into small tubercles.
The surface of the test is granular and very
hard, impregnated with spherical siliceous
bodies. These are 0-30 mm in diameter, have
a granular surface, and are densely packed in
the superficial test, becoming slightly less
dense internally. The test is, however, thin
and very tough throughout. Internally the body
wall is very thin with very strong longitudinal
muscle bands radiating from both siphons and
extending posteriorly to break up into an
irregular network over the gut-loop and pos-
terior end of the body. Circular muscle bands
are thinner but form a more continuous layer
outside the longitudinal bands on the upper
half of the body. The test is invaginated into
the siphonal linings and the same spherical
siliceous spicules are present in this siphonal
lining. There are also minute pointed spines
02 mm long directed toward the aperture.
The dorsal tubercle is large, a U-shaped open-
ing directed toward the left with both horns
turned anteriorly. The branchial tentacles have
primary, secondary and tertiary branches. The
prepharyngeal area is fairly narrow. The dorsal
lamina has short closely-set pointed languets
arising from the border of a narrow membrane.
There are six branchial folds on each side of
the body with 18-21 internal longitudinal
vessels on each fold and two or three vessels
between the folds. The gut loop is embedded
in the body wall and forms a curved open
loop around the posterior end of the body
with arborescent liver tubules rising from the
pyloric region. The gonads on each side of the
body are broken up into about five pairs of
polycarp-like sacs either side of a median
duct. On the left, the gonad is embedded in
the body wall in the primary gut loop; on the
right side of the body the gonad occupies a
corresponding position. The anal border has
six well developed bifid lobes.
Remarks: The most outstanding characteristic
of this species is the very hard "sandpaper-
like" external test, created by the embedded

siliceous spicules. The species may also be
distinguished by the test extensions which
project especially the branchial aperture.

The species appears to be most closely
related to Pyura pachydermatina sub. sp. in-
termedia Michaelsen (which it resembles in
the form of the anal border, the size and
shape of the siphonal spines and in the spheri-
cal siliceous bodies in the test. It may be
distinguished by the absence of the typically
long stalk of P. pachydermatina and by the
shape of the body with the branchial aperture
terminal and not adjacent to the stalk as in
Pyura pachydermatina. The present species may
also be distinguished by the absence of rod-
shaped spicules in the test and by its simple
dorsal tubercular opening which is not con-
voluted as in P. pachydermatina. The body
wall is also less muscular and more closely
adherent to the test, a condition which may
be associated with the very hard and probably
more rigid test.

Pyura scoresbiensis Kott
Pyura scoresbiensis Kott, 1972a, p. 36; 1972b, p. 187.

New Records: Western Port (San Remo)
Distribution: S. Aust: St. Vincent Gulf, Spen-
cer Gulf, Investigator Strait.
Description: Specimens comprise an aggregate
of sandy spherical zooids on long stalks that
taper basally. Both apertures are close together
on the upper surface of the individual. The
body is about 2 cm in diameter. There is an
investment of sand covering the test externally
and the bodies of adjacent individuals adhere
to one another to form the aggregate where
the surface test contacts and adheres to the
sand encrusting an adjacent individual. The
stalks, however, remain free from one another.
There is a slight ridge between the apertures.
There are the usual six branchial folds on
each side of the body. The gut forms a wide
open loop and the rectum extends anteriorly
towards the base of the atrial aperture. Arbore-
scent liver lobes are present in the gastric
region. The gonads consist of approximately
11 pairs of polycarp sacs on either side of a
median duct. They are present in the gut

ASCIDIAN FAUNA OF WESTERN PORT

Ascidia sydneyensis

30— individual showing apertures turned over to-
ward substrate.

Amphicarpa diptycha

31, 32, 33 — variable arrangement of gonads on
right and left sides of the body. 34 — larva.

Cnemidocarpa etheridgii

35 — stalked individual.

Pyura australis

36 — spicules and siphonal spines.

Pyura cataphracta

37— stalked individual. 38 — sessile individual.

39 — spicules and siphonal spines.

Pyura irregularis

40 — siphonal spines.

Pyura albanyensis

41 — siphonal spines.

Pyura stolonifera

42 — siphonal spines. 43 — spicules from body wall.

Microcosmus stolonifera

44 — siphonal spines. 45 — gut loop and gonads.

Microcosmus squamiger

46 — siphonal scales.

PATRICIA KOTT

loop on the left side of the body and in a
corresponding position on the right.
Remarks: The specimens are smaller than
those usually reported for this species. There
is also some variation in body shape which
is spherical rather than the elongate oval as
in the typical specimens described from South
Australian locations. Nevertheless, there is no
character distinguishing them and it is possible
that their small size results from a less favour-
able location possibly at the eastern extent of
the range of this species.

Pyura irregularis (Herdman)
(Fig. 40)

Cynthia irregularis Herdman, 1882, p. 141.
Pyura irregularis; Kott, 1972a, p. 38 and synonymy.
1972b, p. 187.

New Records: Western Port (Crawfish Rock,
8 metres EckJonia holdfasts; Eagle Rock;
Tankerton Jetty); Port Phillip Bay (Morning-
ton Pier).

Distribution: S. Aust.; St. Vincent Gulf; Tas.:
d'Entrecasteaux Channel; Vict.: Port Phillip
Bay; N. S. W.: Port Jackson.
Description: The present specimens were col-
lected in large aggregates together with Micro-
cosmus squamiger. The long siphons are so
oriented that they are able to reach the exterior
despite the other individuals with which they
are so closely aggregated.

Pyura albanycnsis Michaelsen and Hartmeyer

Pyura albanyensis Michaelsen and Hartmeyer, 1928,

p. 435.
Pyura jacatrensis; Kott, 1952, p. 273.
Pyura vittata; Miller. 1960, p. 126; Tokioka, 1952,

p. 134; 1967, p. 202; Kott, 1964, p. 142; 1966, p.

300; 1972a. p. 37; 1972c, p. 243.

New Records: Port Phillip Bay (Mornington)
Distribution: The species has a wide circum-
Australian distribution. It has also been re-
corded from the Palau Is. in the Pacific and
from Ascension Is. in the southern Atlantic.
Description: The test is very tough, leathery
and wrinkled. There are the usual six high
overlapping branchial folds on both sides of
the body. The dorsal tubercle is a large, double
spiral cone completely filling the peritubercular
area. There are needle-like spines lining the
siphons about 01 mm in length. There are

endoearp-like blocks of tissue along the an-
terior limb of the gut loop. The anal border
is bilabiate.

Remarks: This species displays to a very high
degree the taxonomic problems that are en-
countered in the Class Ascidiacca. Species
from many localities and with a wide range
in characters have been included in the syn-
onymy and Van Name (1921, 1945), Kott
(1969, 1972a) and Monniot and Monniot
(1972, 1974) have successively made attempts
to clarify the situation. Briefly, specimens from
the West Indies and the Atlantic, the Indo-
Pacific and the sub- Antarctic (Macquarie,
Kerguelen and Marion Islands) with siphonal
spines variously extending onto the outer wall
of the siphons and the gonads separated into
polycarp-like sacs along cither side of a central
duct have been included in the synonymy.
Monniot and Monniot (1974) have, most
recently, separated the subantarctic form as
Pyura pilosa characterised by the absence of
endocarps on the gut and gonads. In this
species the anal border is lobed. The siphonal
spines are long and one of its most conspicuous
features. Although not described by Monniot
and Monniot these spines must be regarded as
contributing to the diagnosis of the species.

Indonesian and northern Australian species
P. jacatrensis Sluiter, 1890; Hartmeyer,
1919; Michaelsen and Hartmeyer, 1928, were
included in the synonymy of P. vittata by Kott
(1969) but were later (1966, 1972a) ex-
cluded on the grounds that the branchial spines
(003 mm) were very much shorter than those
of sub-Antarctic and Australian specimens as-
signed to this species. The anal border in P.
jacatrensis is simple or very vaguely lobed
and there are no endocarps on gut or gonads
(Monniot and Monniot, 1974).

The West Indies species share, with speci-
mens from Australia {P. jacatrensis; Kott,
1952; P. vittata; Kott, 1972a, c), the Arafura
Sea (P. vittata; Tokioka, 1952), the Palau
Islands (P. vittata Tokioka, 1967) and As-
cension Island (P. vittata; Millar 1960), the
endocarps along the gut loop that are con-
sidered to be characteristic of P. vittata by
Monniot and Monniot (1974). Other speci-

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PATRICIA KOTT

mens from Australia have been re-examined
(Pt. Vernon: Kott, 1966; Moreton Bay:Kott,
1964; Ulladulla, N. S. W.; unpublished and
Port Gregory, Shark Bay, W. A.: unpublished)
and in each case these endocarps are found to
be present. All the Australian specimens
(Kott 1952, 1964, 1966, 1972a, c) and those
from the Arafura Sea and Ascension Island,
have siphonal spines of 01 to 0-275 mm and
these spines confer an irridesccnt greenish
tinge to the siphons of preserved material. In
all cases the anal border is bilabate and plain.
Specimens from Palau Is. and Japan (Tokioka,
1949b, 1950, 1953) also have a generally
plain anal border. However, no endocarps
have been described. The spicules have rot
been described and the siphons are coloured
red. The West Indies species differs from the
Australian and sub-Antarctic forms in the less
conspicuous siphonal spines (for which the
length has not been given: Van Name, 1945
p. 322 "minute short spines, visible only on
some magnification"), the anal border is lobed,
and the siphons are coloured red as in the
Japanese specimens. C. vittata; Oka and C.
karasboja Oka (Oka, 1935) are identical with
the West Indies form in all characters. It is
apparent, therefore, that there are several
species involved in this complex each charac-
terised by a reliably constant assemblage of
characters and in some cases with an over-
lapping geographical range, as set out in the
following Table.

The specimens set out in the synonymy
above conform with the description of P.
albanyensis Michaelsen and Hartmeyer.

Pyura stolonifcra (Heller) subsp.
praeputialis Heller

(Figs. 42-43)

Cynthia stolonifcra Heller, 1878, p. 10,
Cynthia praeputialis Heller, 1878. p. 12.
Pyura stolonifcra f. waia Michaelsen and Hartmeyer,

1928, p. 433.
Pyura stolonifcra: Sluiter, 1927, p. 43; Kott, 1952,

p. 274; MacNae and Kalk, 1958; Kott, 1964, p.

141; Monniot. 1965, p. 100; Day. 1974, p. 35.
Pyura praeputialis; Millar, 1963, p. 738; 1966, p. 372.

New Records; Western Port (Eagle Rock);

Port Phillip Bay (Mornington; Hobsons Bay;

Prince George Light). Ram Head (18 miles

south of Mallacoota, 6 metres).
Distribution: P. stolonijera stolonijera; Cape
Province and Natal (South Africa); Mosam-
bique; Dakar; Morocco. P. stolonijera prae-
putialis; S. W. Aust.; S. Aust.: Outer Harbour;
Vict.; N. S. W.; Qd.: Noosa. The range around
the Australian coast is clearly defined and
accompanied by a decrease in the size of the
individuals at either end of this range (south-
western Australia and Noosa, Queensland).
Description: A single specimen is available
from Mornington and a tight aggregate of
seven individuals from Ram Head. The speci-
mens are of the usual pillar-like form, maxi-
mum height 3-10 cm and maximum diameter,
across the top of each individual, 2-4 cm. In
the specimen from Mornington the posterior
end of the test is produced into irregular
processes for adherence to a rocky substrate.
The specimens have the usual convoluted
double spiral slit on the large hemispherical
dorsal tubercle with the open interval directed
anteriorly. The dorsal lamina has pointed lan-
guets, there are six to seven high overlapping
branchial folds on each side of the body and
the gut forms the usual curved loop termin-
ating in an anus bordered by three shallow
lobes. The liver is large and consists of dense
arborescent tubules. The body musculature is
strong and branches of the longitudinal muscles
from the siphons are inserted into a rim of the
body wall around the anterior surface. These
muscles effect the anterior depression around
the siphons that is characteristic of the species.
Strong longitudinal muscles extend only half-
way down the body. The siphonal spines are
from 0-05 mm to 008 mm long, becoming
larger toward the aperture. Calcareous spicules
(as described by Millar, 1962 for South African
specimens) are present in the ventral part of
the body wall, in the gut and in the endostyle.
Gonads are divided into large paired blocks,
densely arranged along either side of a central
duct, in the gut loop on the left and in a
corresponding position on the right In older
individuals there is a fold of tissue which ex-
tends across the body in front of the atrial
aperture effectively cutting off the posterior
part of the peribranchial cavity as an excurrent

ASCIDIAN FAUNA OF WESTERN PORT

chamber. There is also a dense "fur" of pointed
papillae on the body wall of older specimens
in which gonads are more or less embedded
in the body wall.

Remarks: Kott (1952) has described two en-
vironmental forms of this species one from
estuarine localities and one from open coast
situations. The typical estuarinc form is char-
acterised by the presence of a sandy invest-
ment on the test and is usually short; and
posteriorly the test is produced into a beard
of sandy roots; while the open coast form is
pillar-like owing to the posterior thickening of
the test to form a wide solid gelatinous stand
of the same or of only slightly lesser diameter
than the rest of the body, thus raising the in-
dividual above the substrate. Specimens inter-
mediate between these two forms are known
and the present specimen from Mornington
Pier in which the test is produced into irreg-
ular processes reflects the variability in growth
of the test of this species. Tn the open coast
form the gonads are occasionally broken up
into a single, rather than double, row of sep-
arate sacs. Variations in the leneth of the
branchial tentacles also occur and very small
branchial tentacles have been observed in
estuarine forms. These characters are variable,
however, and cannot be regarded as constant
differences between individuals from these two
environments. The differences in the test espe-
cially appear to result from the individual's
response to different sets of conditions, viz.
the solid gelatinous pillar of test associated
with firm fixation to rocky substrates and to
adjacent individuals where wave action is
strong; the development of long rooting pro-
cesses fixing rounded solitary individuals in
sandy substrates subjected to less turbulent
conditions.

There has been considerable discussion in
the literature on the relationships between the
South African P. stolomfera and the Austral-
ian P. praeputialis. Hartmeyer (1911) sug-
gested that the presence of a seventh branchial
fold was exclusive to South African forms.
There are, however, seven branchial folds on
each side of the body in the present specimens

from Ram Head and this character does not
provide a distinction.

Nor can siphonal spines be used to distin-
guish subspecies. Michaelsen and Hartmeyer
(1928) recognised the subspecies P. stoloni-
fera waia from Western Australia with siphonal
spines 009 to 01 mm long, and P. stolonijera
typica from South Africa and P. stolonijera
praeputialis from New South Wales (Heller's
type specimen) with siphonal spines 02 to
0-024 mm long. The present specimens dem-
onstrate a wide range in length of siphonal
spine between 005 and 0-08 mm while the
spines of typical specimens collected from
Moreton Bay (Queensland) are 0-03 to 0-04
mm long. Spicules (branched) in the branchial
sac of South Africa specimens occur in the
present specimens from Victoria.

Millar (1963) distinguished the South Afri-
can form from the Australian by the absence
of a sunken area around the siphons and by
the posteriorly directed dorsal tubercle open-
ing. Some South African specimens (Millar,
1955) were also distinct in the presence of
four test projections around the apertures.
These differences, however, are not constantly
expressed (Millar, 1966). In fact Cynthia
valdiviae Michaelsen and C. herdmani Von
Drasche (see Michaelsen, 1904), both from
South Africa are identical with the Australian
form in these characters and in others. Day
(1974) has described the depressed area
around the siphons in South African speci-
mens. In Australian specimens, the collar of
test around the siphons is invariably present
when specimens are contracted (i.e. when pre-
served or exposed intertidally). It is probable
that it is not present if individuals are not
contracted. It was not observed by the present
author in large in situ subtidal populations at
Noosa (Queensland) in which the siphons of
all individuals were fully extended in the feed-
ing position. Day (1974) p. 36, has also ob-
served variations in form for the South African
populations that are identical with variations
in the Australian forms, viz: "solitary speci-
mens below tide mark are usually hemispheri-
cal .. . Specimens from S. African estuaries

PATRICIA KOTT

which have root like extensions of the test to
anchor them in the sand".

It is most probable, therefore, that in view
of the general form of the body (and its
capacity to respond to a wide range of en-
vironments) and the similarities between such
features as the dorsal tubercle, the siphonal
spines, and the occasional presence of branched
spicules, that two separate species cannot be
maintained.

Nevertheless, no test projections from the
siphonal region have ever been observed in
the many Australian specimens that have been
examined; and the relationships are most
reliably indicated by regarding the Australian
and South African forms as separate geograp-
hic sub-species, morphologically distinguished
only by a degree of variability of certain
characters in the South African populations,
which are not variable in Australian speci-
mens, i.e. the absence of projections from the
siphonal region (sometimes present in S.
African specimens).

Pyura lcpidoderma Tokioka

Pyura lcpidoderma Tokioka, 1949, p. 10. Kott, 1966,
p. 299.

New Records; Port Phillip Bay.
Distribution: Qd.: Hervey Bay. Japan.
Description: A single flattened specimen, basal
diameter 1 cm, heighth 2 mm is available, fixed
to a Mytilus shell. The siphons are on the
upper surface and are almost sessile. The sur-
face layer of test is marked off into polygonal
scale-like thickenings that are most conspic-
uous around the apertures where they em-
phasise furrows along the short siphons. The
body musculature consists of longitudinal bands
from both apertures which cross one another
on the body. There arc six branchial folds
on each side of the body with closely set in-
ternal longitudinal vessels. The liver branches
off the gut in the pyloric region and divides
into arborescent branches along its length.
The gut loop is narrow and the anal border
has shallow rounded lobes, the rectum turns
anteriorly and is very short. The gonads, in
the gut loop on the left and in a corresponding
position on the right, consist of polycarp-like

sacs along either side of common ducts
opening at the base of the atrial aperture with
the anus.

Remarks: Attention has already been drawn
(Kott 1966, 1969) to the similarity between
this species and the Antarctic P. squamata
Hartmeyer (see Kott 1969, p. 136). The oc-
currence of the species in Port Phillip does
to some extent satisfy the discontinuity in the
records of the antarctic species and the north-
eastern Australian Japanese species respec-
tively. It is possible that further collecting will
demonstrate that their ranges are continuous
and that only a single species is represented.

Halocynthia hispida (Herdman)

Cynthia hispida Herdman, 1881, p. 61.
Halocynthia hispida; Kott, 1968, p. 77 and synonvmy;
1972b, p. 189.

New Records: Western Port (Crawfish Rock;
some on Ecklonia holdfasts; Eagle Rock).
Distribution: S. Aust.: St. Vincent Gulf; Tas.:
Bass Strait, d'Entrecasteaux Channel, Maria
Island; N. S. W.: Port Jackson. Ceylon; and
off the west coast of North America and
Japan, (see Kott, 1968).
Description: The present specimens demon-
strate the range from the individuals covered
with branched test tubercles to those in which
there are no tuberculous extensions of the
test, (see Kott, 1968).

Herdmania inomus (Savigny)

Cynthia mom us Savigny, 1816, p. 143.
Herdmania momus; Kott, 1972b, p. 189; 1972c, p.
255 and synonymy.

New Records: Western Port (Flinders Jetty,
Eagle Rock).

Distribution: N. W. Aust. (Broome); S. W.
Aust.; S. Aust.; Vict; N. S. W.; Qd., Arafura
Sea; Indonesia; Fiji; the Palau Islands; Tahi-
ti; Japan; the Indian Ocean; the Red Sea;
South Africa; and the West Indies. Kott (1972)
has drawn attention to the lack of distinction
between the Indo-Pacific populations of this
species and those from the Atlantic, H. momus
pallida (see Van Name, 1945). It is most
probable, therefore, that H, momus is a cir-
cumtropical species extending into temperate
regions in the southern parts of its range, i.e.

ASCIDIAN FAUNA OF WESTERN PORT

around the South African and south Aus-
tralian coasts.

Description: The specimen from Flinders jetty
is 20 cm long and completely invested with a
colony of Didemnum patulum, leaving only
the apertures free.

Microcosmus australis Herdman

Microcosmus australis Herdman, 1889, p. 23. Millar,
1963, p. 741; 1966, p. 373. Kott, 1972e, p. 53.

Microcosmus claudicans sub sp. australis; Michaelsen
and Hartmeyer, 1928, p. 404 and synonymy. Kott,
1952, p. 288.

ICynthia solanoides Herdman, 1899, p. 29.

IMicrocosmus solanoides; Kott, 1952, p. 289.

New Records: Western Port (Crawfish Rock).
Distribution: see Kott 1972e
Remarks: There seems little to distinguish this
species from M. solanoides Herdman except
the siphonal denticles which Kott (1952) des-
cribed as not curved and much smaller than
those of the present species. Herdman's type
specimen from Port Jackson is the only record
of M. solanoides and it is likely that it cannot
be separated from M. australis.

Microcosmus nichollsi Kott

Microcosmus nichollsi Kott, 1952, p. 290; 1972c,
p. 245 and synonymy.

New Records: Western Port (Crawfish Rock,
8 metres, on Ecklonia holdfasts).
Distribution: S. Aust.: St. Vincent Gulf; Vict:
Flinders.

Description: Only a single specimen is avail-
able, 1 cm long. The surface is sandy and
both apertures are anterior and almost sessile.
The usual pockets or valves are present at
the base of the atrial siphon, and the usual
spines and scales are present in the siphonal
lining. There are seven branchial folds on each
side of the body with one or two internal
longitudinal vessels between the folds. The
gonads are separated into three blocks and on
the left the most proximal section of the gonad
crosses into the pole of the gut loop.

Microcosmus helleri Herdman

Microcosmus helleri Herdman, 1881, p. 54 Van Name,
1945, p. 349 and synonymy; Kott, 1972e, p. 54
and synonymy.
New Records: Western Port (Crawfish Rock,
among Ecklonia holdfasts; Eagle Rock).

Distribution: W. Aust.: Cape Jaubert to Fre-
mantle; S. Aust.: St. Vincent Gulf; Qd.: Great
Barrier Reef, Gulf of Carpentaria, Torres
Strait; Malaysia. Portugese East Africa
(Michaelsen 1918). West Indies (Van Name
1945).

Description: The individuals are upright and
more or less egg-shaped with a terminal
branchial aperture and the atrial aperture about
half way down the dorsal surface. Around
each opening the test is produced into lobes
which fold over when the aperture is con-
tracted. Superficially the test is covered with
long branched hairs that are obscured by a
coating of sand which they enmesh. Beneath
this sandy coating the test is very thin and
brittle. There are four, very strong, tongue-like
projections into the cavity of the branchial
siphon at its base. There are three pockets
at the base of the branchial siphon, formed by
folds of the siphonal lining and these undoub-
tedly act as cuspid valves. There are six
branchial folds on each side of the body. The
dorsal tubercle is U-shaped with both horns
turned in and completely fills a fairly shallow
peritubercular area. The gut forms the usual
narrow loop and in the pyloric region there
are dense parallel glandular folds or lamellae,
joined together by an external membrane, re-
presenting the liver. The long gonad on each
side of the body is divided into three separate
sections, and on the left crosses the intestine
into the pole of the gut loop.
Remarks: The hard cartilage-like projections
at the base of the siphon together with relat-
ively small number of wide overlapping branch-
ial folds characterise this species. The test
lobes around the apertures are sometimes
simple, but often are well developed, tuber-
culous or branched. In some specimens there
is no coating of sand but externally the test
is very hard and produced into pointed and
sometimes branched papillae.

Microcosmus stolonifera Kott

(Figs. 44, 45)

Microcosmus stolonifera Kott, 1952, p. 291; 1972c,

p. 245 and synonymy.
New Records: Western Port (Crawfish Rock).

PATRICIA KOTT

Distribution: Previously recorded from QUI.
(Moreton Bay) S. Aust. (St. Vincent Gulf)
and Tas. (Tiny Is.).

Description: Posteriorly the test of these in-
dividuals is produced into an irregular root-
like structure, sometimes long branched and
sturdy, sometimes there is a double projection.
Occasionally individuals are aggregated to-
gether. Both apertures are depressed into the
upper surface and surrounded by a raised,
rounded fold of test. The test is very hard,
thin and stiff with a dense layer of embedded
sand. There are overlapping curved spines
lining the branchial siphon 06 mm long on
their concave side but extending from a long
base of about the same length. Short, irregular
languets are sometimes present on the pre-
pharyngeal band. The branchial tentacles have
primary, secondary and minute tertiary
branches. The dorsal tubercle is relatively
small in the centre of the peritubercular area
and has a simple, U-shapcd opening, some-
times with a single horn turned in. There is
a long, smooth-edged dorsal lamina. There
are seven high and overlapping branchial folds
on each side of the body, with 15-20 internal
longitudinal vessels per fold and six to eight
stigmata per mesh. The gut forms a narrow
loop with liver lamellae in the pyloric region.
Single rounded or sometimes irregular gonads
are present on each side of the body. The
gonad on the left is in the secondary gut loop
and does not extend into the primary gut
loop.

Remarks: The rounded fold of test around the
upper surface enclosing the apertures is not
always present and depends to some extent
on the size of the individual. The species is
distinctive, however, in the presence of the
gonad outside the primary gut loop and in
the size and form of the siphonal spines to-
gether with the high overlapping folds in the
branchial sac.

Microcosmus squamiger Michaelsen
(Fig. 46)

Microcosmus claudicans sub. s. squamiger Michaelsen,
1928, p. 405.

Microcosmus squamiger; Kott, 1972a, p. 43 and syno-
nymy.

New Records: Western Port (Crawfish Rock;
Eagle Rock; Rutherford Channel). Port Phil-
lip Bay (Mornington Pier, 9 to 35 metres,
dense rock, vertical and horizontal clumps
fixed to oyster shells; Williamstown, 5 metres,
common on rocks).

Distribution: W. Aust: Shark Bay to Albany;
S. Aust: St. Vincent Gulf; N.S.W.: Port Jack-
son; Qd.: Bowen and Rockhamton. Red Sea.
Undoubetdly there has been great confusion
between this and related species (see Michael-
sen and Hartmeyer, 1928) and it is probable
that there is a wider distribution in the Indian
Ocean,

Description: Rounded individuals are present
in sometimes very large aggregates. The aper-
tures are about one third of the body length
apart and sessile. The animals are whitish
to pinkish-brown, sometimes smooth, occasion-
ally with some embedded sand. The surface
is often very uneven and wrinkled, especially
around the siphons. There are minute over-
lapping siphonal scales about 001 mm long.
There are eight to nine overlapping branchial
folds on each side of the body. The dorsal
tubercle is always a double spiral cone, the
gut forms a narrow loop, the gonads are
separated into three sections and cross into
the primary gut loop.

Remarks: This is a common species in those
areas from which it has been recorded and
is distinguished by the very small overlapping
siphonal scales, the dorsal tubercle, the
leathery test and the absence of a dense,
sandy coating and the large number of over-
lapping branchial folds.

Molgula sabulosa Quoy and Gaimard

Ascidia sabulosa Quoy and Gaimard, 1834, p. 613.
Molgula sabulosa; Michaelsen and Hartmeyer, 1928,

p. 449 and synonymy. Kott, 1972c, p. 248 and

synonymy.

New Records: Western Port (Crawfish Rock,
San Remo; Shoreham). Port Phillip Bay.
Distribution: Known only from Albany, (W.
Aust.) and Port Phillip Bay (Vict.)
Description: Large spherical specimens with
characteristic hollow test lobes formed around
the apertures. The individuals often form

ASCIDIAN FAUNA OF WESTERN PORT

aggregates. The test is thin but stiff with
adherent sand. The apertures are close to-
gether on the upper surface but are not in
a depressed pit as in M . mollis, and do not
have a thickened ridge of test extending along
the dorsal line between them. Each aperture
is surrounded by a rosette of hollow rounded
lips of the test, 6 around the branchial aper-
ture and four around the atrial aperture.
These fold inwards over the aperture and
each aperture together with its surrounding
test projections, occupies a circular depres-
sion in the upper surface of the body. The
body wall projects into these hollow lobes.
The dorsal tubercle occupies the right hand
side of the peritubercular area and there is
elongate ganglion slightly to the left. The
dorsal lamina is rather long. There are seven
narrow branchial folds on each side of the
body with only three internal longitudinal
vessels, one on the edge of the fold and the
other two ventrally. Stigmata are irregular
between the folds and between the dorsal
lamina and the first fold. The infundibula are
tightly coiled and bifurcate in the summit of
each fold. The gut forms a long narrow curved
loop and the plain bordered anal opening
is at the base of the atrial siphon. There
are many fine longitudinal liver lamellae at
the cardiac end of the stomach but at the
pyloric end there are minute arborescent
lobes. The kidney is long and curved on the
right hand side of the body and the long
tubular ovary extends parallel to its dorsal
surface. Testes follicles are long narrow and
deeply lobed at their outer edge tapering to
the vasa efferentia at the end of the ovarian
tube where they join into the short vas def-
erens that extends onto the mesial surface
of the ovarian tube and opens to the peri-
branchial cavity there.

Remarks: This species is distinguished from
M. mollis by the short vas deferens
(which in some specimens is turned antero-
ventrally), by the longer dorsal lamina and

by the hollow lobes of test that protect the
apertures.

Mogula mollis Herdman

Mogula mollis Herdman, 1899, p. 54; Kott, 1952,
p. 298; 1964, p. 144; 1972a, p. 45 and synonymy.
Mogula sabulosa; Kott, 1972a, p. 190, 1972d, p.

248.

New Records; Western Port (Crawfish Rock).
Distribution: S. Aust: St. Vincent Gulf; east-
ern Australia to Indonesia.
Description: The specimens are smaller than
is usual for M. sabulosa and are laterally
flattened and covered with delicate hairs to
which sand adheres. There are small test pro-
jections around the apertures but these are
not hollow as in M. sabulosa. The apertures
are close together on the upper surface of
the body both sunk deeply into the surface,
with a ridge of slightly thicker test between
them along the dorsal line. The apertures
are directed away from one another. The
body wall has very strong muscles in the
siphons and around the anterior part of the
body at the base of the siphons but poster-
iorly the musculature is weaker. The branch-
ial folds are broad, overlapping and deeply
curved and the dorsal lamina is very short.
There may be as few as 4 internal longitud-
inal vessels on both sides of each fold. The
gut forms a long, very deeply curved loop
which enclosed the gonad on the left side of
the body. At the cardiac end of the gastric
region there are short transversely oriented
liver lamellae but at the pyloric end there
are longer and longitudinally arranged diver-
ticulae. The testis follicles are grouped around
the proximal end of the ovarian tube. They
are long narrow and wedged shaped in out-
line, converging to paired ducts which join
into a long vas deferens that extends along
the middle of the mesial surface of the
ovarian tube and opens at the base of the
oviduct. On the right side of the body the
testis lobes are accommodated in the curve
of the kidney and not anterior to it as in
M. sabulosa.

PATRICIA KOTT

I»RW uvtr

niii»»sTQim_

47 — Map of region.

ASCIDIAN FAUNA OF WESTERN PORT

CHARACTERISTICS OF THE ASCIDIAN FAUNA
(Fig. 48)

Most collecting of the Victorian ascidian
fauna has been done in either Western Port
or Port Phillip Bay. It is probable that a
proportion of the very large number of spec-
ies recorded from these two locations also
occur on the open coast where a relatively
limited range of habitats has been sampled.

The material reported on by Millar (1966)
from Port Phillip Bay was collected by bot-
tom sampler and the species taken represent
the fauna of the sea floor. Kott (1952, 1957,
1962, 1963) reported largely on intertidal
collections; while the present mainly subtidal
collections have been taken manually from
rock ledges, caves and from the sea floor
and represent the larger components of the
benthic fauna.

There are 26 species that have been re-
corded from Port Phillip Bay but do not
occur in Western Port. Three are probably
introduced (viz. Ciona intestinalis; Ascid-
iella aspersa; Kott, 1952; and Styela clava:
Holmes, 1976). Ritterella assymmetrica
Millar, 1966 and Protopolyclinum sabulosa
(Millar, 1963) are endemic to Port Phillip
Bay. Of the remaining 21 species, 12 are
within their Australian range; and 7 are tem-
perate species at the eastern or northern ex-
tremity of their range (including Polycitorella
mariae; Millar, 1963, previously recorded
from South Africa and New Zealand). For
only three of these species does their occur-
rence in Port Phillip represent the southern
limit of their range on the eastern coast of
Australia.

There are 45 species that have been re-
corded from Western Port but have not been
taken from Port Phillip. Twenty four of
these are recorded only from Crawfish Rock
and at no other location and a further 10
occur only at the adjacent Eagle Rock or at
both Eagle Rock and Crawfish Rock. Of
this 34 species one is endemic (Polysyncraton
victoriensis sp. nov.); Dumus arenijerus pre-
viously known only from New Zealand is re-
corded from Australia for the first time; 16
are in the middle of their range; eight are

at the southern limit of their range and eight
are at the eastern limit of their range. Of
the species that are also recorded from other
parts of Western Port three are in the middle
of their range, five are at the eastern end of
their range and three are at the southern end
of their range (Fig. 48; Tables 1, 2).

Comparison of the geographical affinities of
ascidian fauna of Western Port and Port Phillip
Bay

Introduced Species

Endemic Species

rum

Species at southern
limit of range

F=l

Species at eastern
limit of range

1 1

Middle of range

PP Port Phillip Bay

WP Western Port

CR C

rawfish Rock

(Western Port)

ER E

agle Rock

(Western Port)

(WP) Western Port but not

from CR or ER

Scale

CZD

= 1 species

PP*WP PP CfUER (WP)

48 — Histograms showing geographic affinities of
the Western Port and Port Phillip Bay ascid-
ian fauna.

TABLE 1

Geographical affinities of ascidian species from
Port Phillip Bay*

Species

Geographical
affinity

Ciona intestinalis
Protopolyclinum sabulosa (Millar,

1963)
Polycitorella mariae', Millar, 1963
Distaplia viridis; Kott, 1972a
Distaplia stylifera; Kott, 1972b

Introduced

Eastern limit
S. Africa; N.Z.
Eastern limit
Middle of range

PATRICIA KOTT

Middle of range

Endemic

Middle of range
Southern limit
Eastern limit
Middle of range
Middle of range
Middle of range
Introduced
Middle of range
Eastern limit

Middle of range
Southern limit

Middle of range
Middle of range
Middle of range
Introduced
Eastern limit
Northern end of

range
Middle of range
Southern limit

* A recent reference is given for each species not
discussed above.

TABLE 2

Cystodytes dellechiajei; Kott, 1972b
Ritterella assymmetrica Millar,

1966
Euherdmania australis\ Kott, 1972b
Aplidiwn solidum; Millar, 1963
Didemnum lambitum
Perophora hutchisoni; Millar, 1966
Ascidia gemmata
Ascidia aclara; Millar 1963
Ascidiella aspersa; Millar 1966
Corella eumyota; Knott, 1972a
Oculinaria australis; Millar, 1966
Botrylloides magnicoecus; Millar,

1966
Polyandrocarpa lapidosa
Polycarpa pedimculata; Millar,

1966
Cnemidocarpa etheridgii
Styela plicata; Millar, 1966
Styela clava; Holmes, 1976
Astereocarpa cerea; Millar, 1966
Pyura fissa; Millar, 1966

Pyura albanyensis
Pyura lepidoderma

Geographical affinities of ascidian species from Western Port 11

c o

.2 °-

Species

o <u

1—1 crt

Geographical affinities

i-H

Podoclavella cylindrica

Eastern limit

Oxycorynia pseudobaudinensis n. sp.

Eastern limit

Eudistoma pyri forme

Middle of range

Polycitor giganteum; Kott,

1972b

Middle of range

Atapozoa mirabilis

Eastern limit

Sycozoa cerebriformis

Middle of range

Sycozoa pedunculata

Middle of range

Pseudodistoma cereum

Middle of range

Dumus areniferus

Single Australian record

Polyclinum marsupiale

Middle of range

Aplidium australiensis Kott

, 1963

Eastern limit

Aplidium parvum Kott, 1963

Eastern limit

Aplidium pliciferum; Kott,

1972a

Middle of range

Aplidium depressum

Southern limit

Aplidium lobatum

Southern limit

Aplidium triggiensis

Eastern limit

Aplidium opacum Kott, 1963

Middle of range

Synoicium hypurgon

Middle of range

Sidneyoides tamaramae

Southern Limit

Didemnum moselyi

Middle of range

Didemnum patulum

Middle of range

Didemnum turritum

Eastern limit

Didemnum august i

Eastern limit

Didemnum roberti

Eastern limit

Didemnum spongioides

Middle of range

ASCIDIAN FAUNA OF WESTERN PORT

Species

Didemnum skeatii
Didemnum candidum
Trididemnum cerebriforme
Trididemnum cy clops
Lissoclinum fragile
Lissoclinum ostrearium
Diplosoma translucida
Diplosoma rayneri
Polysyncraton orbiculum
Polysyncraton victoriensis n. sp.
Phallusia depressiuscula
Ascidia sydneyensis
Symplegma viride
Amphicarpa diptycha
Botrylloides nigrum
Botrylloides leachii

Polycarpa thely panes

Pyura australis

Pyura cataphracta

Pyura irregularis

Pyura scoresbiensis

Pyura stolonijera praeputialis

Halocynthia hispida

Herdmania momus

Microcosmus australis

Microcosmus nichollsi

Microcosmus helleri

Microcosmus stolonijera

Microcosmus squamiger

Molgula mollis

Molgula sabulosa

a o

.2^

M
o

Geographical affinities

a -

5.S

Southern limit

Middle of range

Southern limit

Southern limit
Southern limit

Middle of range

Eastern limit

Endemic

Middle of range

Eastern limit

Middle of range

Middle of range
Southern limit

Eastern limit

Southern limit

X
X

Middle of range
Eastern limit

Middle of range

X
X

Middle of range
Middle of range

Southern limit

Eastern limit

Middle of range

X
X

Southern limit
Eastern limit

* The most recent reference is given for each species not discussed above.

It is probable that both Port Phillip Bay
and Western Port provide different habitats
for ascidian species since only a small group
of species (the majority of these well within
the limits of their geographic range) have
sufficiently unrestricted habitat requirements
to be present in both locations. Larger num-
bers of species are recorded from either Port
Phillip Bay or Western Port but not from
both. The most striking differences in the
biogeographic affinities of the ascidian fauna
at these two locations is the high diversity of
species and the high percentage of northern
forms at the southern limits of their range
that occur in Western Port. These differences
are not associated with a random distribution
of habitats since the majority of species have

been taken at Crawfish Rock and also at
adjacent Eagle Rock and have not been re-
corded at more southerly locations in West-
ern Port. Both these stations are in the north-
ern section of the bay, where there is comp-
lete protection from oceanic swell and where
extensive tidal flats draining into the main
channel may modify the temperate marine
environment. This may contribute to the
diversity of environmental conditions that
support such a diverse fauna with such a
high proportion of northern species.

The most diverse assemblage of species
has been taken from Crawfish Rock, on a
30° slope of soft brown coral and occasional
sandstone boulders. The species are largely
encrusting aplousobranch forms with vivi-

PATRICIA KOTT

parous larvae but there are also small leathery
oviparous stolidobranch species that produce
root-like structures and form aggregates. The
stalked species Pyura australis only rarely
occurs here, and large stolidobranch and
phlebobranch forms (Phallusia depressiuscula,
Herdmania momus, Cnemidocarpa etheridgii,
etc.) that require smooth and firm surfaces
for fixation are not present.

The turbidity of the water and its effect
on the light intensity at greater depths does
not appear to limit ascidian distribution at
this location.

Similar, though not such dense nor diverse
associations of species occur at Eagle Rock
and in the Rutherford Channel.

It is possible that the less diverse fauna of
Port Phillip Bay is a result of environmental
disturbance. The presence there of (probably)
introduced species Ciona intestinalis, Ascidi-
ella aspersa and Styela clava (see Holmes,
1976) may be evidence of this disturbance.

The occurrence of Dumus areniferus B re-
win, previously known only from New Zea-
land, increases the number of temperate spec-
ies that are known to occur in both southern
Australian and New Zealand waters (Kott,
1974). It is unlikely to have been introduced
on ship's hulls and its Australian occurrence
may have previously been overlooked.

STATION LISTS

WESTERN PORT

CRAWFISH ROCKS, tidal currents, 5 knots
? metres:

Aplidium triggiensis

Trididemnum cerebriforme
Intertidal:

Microcosmus squamiger

Microcosmus australis
0-15 metres:

Sycozoa cerebriformis

Dumus areniferus

Aplidum lobatum

Sidneoides tamaramae

Didemnum skeati

Didemnum spongioides

Amphicarpa diptycha

Pyura irregularis

Halocynthia hispida

Mogula mollis
8 metres, Ecklonia 'holdfasts':

Aplidium lobatum

Didemnum candidum

Didemnum mosleyi

Didemnum patulum

Polysyncraton victoriensis

Botrylloides leachii

Pyura australis

Pyura cataphracta

Pyura irregularis

Halocynthia hispida

Microcosmus helleri

Microcosmus nichollsi

Microcosmus stolonifera

Microcosmus squamiger
12 to 24 metres:

Oxycorynia pseudobaudinensis

Sycozoa cerebriformis

Eudistoma pyriforme

Polycitor giganteum

Pseudodistoma cereum

Synoicium hypurgon

Polyclinum marsupiale

Aplidium lobatum

A plidium depressum

Didemnum moseleyi

Didemnum patulum

Didemnum turritum

Didemnum augusti

Didemnum skeati

Didemnum spongioides

Didemnum roberti

Polysyncraton orbiculum

Lissoclinum ostrearium

Diplosoma rayneri

Ascidia Sydney ensis

Botrylloides nigrum

Symplegma viride

Amphicarpa dytycha

Pyura australis

Halocynthia hispida

Microcosmus helleri

Microcosmus stolonifera

Microcosmus squamiger
13-26 metres:

Amphicarpa dipytcha

Pyura australis

EAGLE ROCK, 15 metres:

Trididemnum cyclops
Didemnum mosleyi
Didemnum patulum
Didemnum turritim
Didemnum roberti
Didemnum skeati
Lissoclinum fragile
Diplosoma translucidum
Ascidia sydneyensis
Botrylloides nigrum
Pyura irregularis
Pyura stolonifera praeptialis
Halocynthia hispida
Herdmania momus
Microcosmus helleri
Microcosmus squamiger

RUTHERFORD CHANNEL, fast current, 5 metres:
Sycozoa penduculata
Aplidium depressum
Microcosmus squamiger

ASCIDTAN FAUNA OF WESTERN PORT

TANKERTON JETTY, 7 metres:
Polycitor gigan/eum
Atapozoa mirabilis
Sycozoa cerebriformis
Aplidium pliciferum
Phalhtsia depressiuscula
Pyura irregularis

SHOREHAM:

Ascidia syneyensis
Molgula sabulosa
SAN REMO:

Pyura scoresbiensis
Molgula sabulosa

BALNARRING BEACH:

Podoclavella cylindrica
FLINDERS JETTY, 3 metres:

Oxycorynia pseudobaudine nsis
Podoclavella cylindrica
Trididemnum cyclops
Lissoclinum ostrearium
Phallusia depressiuscula
Ascidia sydneyensis
Polycarpa thelypanes
Herdmania momus

PORT PHILLIP BAY

WILLIAMSTOWN, 5 metres (common on rocks):
Ascidia sydneyensis
Mierocosmus squamiger

HOBSONS BAY, 13 metres:

Didemnum lambitum
Ascidia sydneyensis
Phallusia depressiuscula

MORNINGTON, 12/10/69, bottom patchy, rock and
sand, coll. Kevin Duke, 1-2 metres:
Pyura albanyensis
Pyura stolonifera praeputialis
MORNINGTON PIER, 12/10/69, sparse rock, calm
waters, coll. Kevin Duke:
Ascidia gemmata
Cnemidocarpa etheridgu
MORNINGTON PIER, 12/10/69, dense rock, coll.
Kevin Duke, 8-11 metres:
Pyura irregularis
Mierocosmus squamiger
YARRA RIVER, 0-3 metres, 28/4/72, No. 3 Oil
Wharf, Coll. J. E. Watson.
Ciona intestinalis
PORTSEA, 15/9/1957:

Amphicarpa diptycha
MORDIALLOC BEACH, Nov, 1888, Coll. W.K.;

Aplidium pliciferum
Botrylloides nigrum
ZOOBENTHOS SURVEY, Fisheries and Wildlife De-
partment, Coll. G. Poore: Off BRIGHTON,
21/10/69, Station 906, sandy bottom, 10 metres;
middle of northern part of Gulf, 10/2/69,
Station 915, silty-clay bottom, 19 metres; of!
GEELONG, 12/2/70, Station 940, silty clay
bottom, 8 metres:

Ascidia sydneyensis
South of POINT WILSON, 12/2/70, Station
942, silty clay bottom, 7 metres:

Ascidia sydneyensis

Molgula sabulosa
North-west of PORT ARLINGTON, 11/2/70,
Station 930, silty clay-sand bottom, 10 metres:

Ascidia gemmata
North of PORT ARLINGTON, 18/2/71, Station
931, silty sand-shell bottom, 15 metres; north-
west of ROSEBUD, Station 982, sandy bottom,
18 metres:

Sycozoa pedunculata

Cnemidocarpa etheridgii
East of ST. LEONARDS, 16/2/7 1, Station 960,
sandy bottom, 10 metres:

Moluga mollis

Off BRIGHTON, 9/3/71, Station 1218, sandy
bottom, 4 metres:

Pyura lepidoderma

Pyura stolonifera praeputialis
Off BRIGHTON, 9/3/71, Station 1226, silty-
clay bottom, 8 metres; half-way down eastern
shore of the Bay, 10/3/71, Station 1241, sandy
bottom; south of SOUTH WERRIBEE, 11/3/71,
Station 1252, sand-gravel bottom, 4 metres:

Pyura stolonifera praeputialis
PORT PHILLIP SURVEY:

area 5, POPES EYE, 15/5/63:

Polyandrocarpa lapidosa
area 30, PRINCE GEORGE LIGHT:

Pyura Stolonifera praeputialis
ARTIFICIAL REEF:

Ciona intestinalis

Sycozoa pedunculata

Ascidia syneyensis

Botrylloides nigrum

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MORTUARY CUSTOMS OF NORTHEAST ARNHEM LAND: AN
ACCOUNT COMPILED FROM DONALD THOMSON'S FIELDNOTES

By Nicolas Peterson 1

Dept. of Prehistory & Anthropology, Australian National University.

Tf a man could but follow all that takes place when a yarkomrri [important] man dies he
would understand almost all of the culture of these people.' — Fieldnotes, July 29, 1937.

On learning of a man's death close female
relatives throw themselves on the ground and
hit their heads with knives, bone points or
sticks, until blood flows. Some close male rela-
tives may weep and the son of the dead man is
likely to become angry and aggressive towards
his father's enemies. Like the son, the actual
sister's son may also become angry, recalling
past quarrels in which his mother's brother had
figured, regardless of whether he had been in
the right or wrong. Distant male relatives in the
camp at the time of death sit quietly with
bowed heads.

The wife of a deceased man usually sits
down beside him, places the head in her lap
and with her left arm around the body cries
all day. At night she may lie beside him sur-
rounded by other camp members who weep and
sing through the night. The songs indicate the
path the deceased's spirit, birrimbir, should
take.

If people bring news of a death to a camp
they do not announce the name of the person
but only mention that somebody has died. A
senior man in the group then sings a song
formally announcing the death and at the end
of the song identifies the person and indicates
the cause of death without using their name.
BODY PAINTING
Most deceased people are painted with a clan
design, mlntjL The design painted on the body
should be and usually is that belonging to the
person's own clan. However, absence of the
right people may mean that the clan design of
the actual MM clan is used. Once painted the
design must not be seen by women or children
so the painting is often carried out at the edge
of the camp and when the painting is completed
the corpse is covered with paperbark, only the
face being left bare. Young men who see a clan

design for the first time have underarm sweat,
bunggan wurdoi, of an older man, rubbed over
their eyes.

The body is first rubbed with red ochre and
then painted by one or two men, preferably
of the opposite moiety to the deceased, who
are good hands at painting, kong mintjimirri
The most frequently chosen relatives are from
the categories FZS, MBS, ZS, MB and MF/
FMB. If the painters are of the same moiety
they are likely to be close WMB, ZDS, or FZDS
but never actual F, B or S. Whoever they are,
they are referred to as the kong wukundi, 2
hands tabu from death, and after they have
completed the painting coat their arms and
hands with red ochre, eat apart for about a
week, refrain from sexual intercourse and do
not go near water. They lived with their wives
some distance from others in the camp. The
kong wukundi and their wives cook all their
food in a sand sculpture (wandjur — discussed
below) and put all their food scraps into
another.

Case 1. Bodv painting (see Plate 4). An old Obul-
karra [Wulkara of Warner, see 1958: 46] woman
died at Milingimbi on Sept. 19, 1935. She had
been declining for some months and was very thin
from the effects of leprosy. As she was old there
was very little crying, but Thomson was suprised
at the apparent indifference of all the imme-
diate relatives. The day following the death a
Wunguri clansman who called the deceased muk-
kulmal (FZ) and a Tjambarapoingo man who
called her momalkor (WMM) painted the body.
The painting was carried out under the shade
of a big tree about fifty yards from the camp. The
husband and a few other men came to and from
the place at which the body was painted from
time to time. Ordinarily her husband would have
assisted in the painting but he was too old and
could not see properly. The brothers of the de-
ceased, as is always the case, could not touch the
body. At the conclusion of the painting the kong
wukundi painted their hands and arms below the
elbow with red ochre and that night held a mant-
jarr ceremony. This took place on the fringe of

NICOLAS PETERSON

the camp. A sand sculpture, wandjur (see below)
representing the clan well of the deceased was
moulded on the ground. The sculpture was associa-
ted with the edible corm of Eleocharis dulcis, called
rakai, but the significance of this was hard for
Thomson to follow at the time [as he had only
just arrived in Arnhem Land]. The men sat around
the sculpture singing for an hour or two, and then
the women danced behind them. A fire was lit
in the centre (manotji [literally eye but also used
of a certain kind of well]) of the sand sculpture.
Leaves were then heated in the fire by a man
whose mother came from the 'Wulkarra [that is
Obulkarra] side'. All the immediate relatives gath-
ered inside the sculpture about the manotji, and
a 'big' man dalkarramirri, called out in loud voice:
'KuritaF (fire).

'Ye'h- replied the participants, with a long his-
sing shout.
Fire was called because the ancestral woman,
mialk kortjurino, burned the grass to clear the
ground while looking for rakai.

The participants then shouted 'Wap wap! wap!',
the sound of the fire burning.

The dalkarramirri then called out a series of
big names with the participants answering 'Ye'h!'
to each.

'NunimarraV (big name [of fire?]).
'Baltjau'wuma' (a big light made by fire flaring

up in thick grass).
'Matauwupuptum' (leaping tongues of fire).
'Birraudun' (cleansed by burning).
Throughout the calling leaves were being heated
in the fire and used to strike the bodies of the
people standing in the sculpture. After use they
were burnt in the fire and to the accompaniment
of singing, smothered with earth, 'that mialk
[woman] walk about now — come along clean place'.
Then the men sang of marratnatta, a rodent
that followed the fire and established itself in the
plains after the burning; and then of dog (workan)
who smelt the rats; and after that of plain cock-
atoo (Corella sp.) kai karra.

The ancestral woman now looked about for
the rakai. Next the men sang of mist (kardany)
which is like fine smoke; then of a spider making
its [nest/webb?] in the damp grass. The last song
was about the wind that follows the time of burn-
ing the countryside.

Close relatives are not free from all tabu until
after a second cleansing ceremony.

HAIR

Before burial of the body all the head hair
is pulled out by the kong wukundi. The beard
is also pulled out with the aid of hot bees wax.
The hair is kept in a basket and sometime
during the following weeks is sent by the kong
wukundi to a fairly distant relative of either
moiety to turn into a string decorated with
feathers, marngarai (Kopapoingo, Tjambara-
poingo and Koiyamillilli) or yiritpal (Wunguri).
The maker of the yiritpal or marngarai rubs
red ochre on the hair as soon as he receives it

MEM. NAT. MUS. VICT. 37 PLATE 4

An Obulkarra woman being painted with a clan
design after death (Case 1).

MORTUARY CUSTOMS

and is given a present of vegetable food from
the marramorkoimirri (the deceased's patri-
clansmen). Usually the maker is of the opposite
moiety to the deceased but he does not have to
be; common choices are people in the category
of MF/FMB, MB, MBS, WMMB. One to two
years later it will be completed and returned
to the close kinsmen of the deceased eventually
being given to his son, if adult. A large presen-
tation is made to the maker; this formerly
included vegetable food, cycad bread and
hooked and short spears. The string is then
used as a belt to be worn in ngarra ceremonies
and during fights and makaratta peace-making
settlements. Eventually the string is cut up to
form the 'arms' of men's sacred baskets.

BURIAL

Either the same day as the painting or the
day afterwards the body is buried. There are
two types of burials: either in the ground or
in a tree.

A grave, molo (referring specifically to the
heap of earth covering the grave) is usually
1 m deep and long enough for the corpse to
lie extended, on its front. There seems to be
some variation of opinion as to which way a
deceased man's head should point. In a dis-
cussion on the matter a Djinang and a Lia-
gallauwumirri man maintained it should be
towards the clan well while a Kopapoingo and
Tjambarapoingo man maintained it should be
eastwards.

If a body in a grave is covered with heavy
logs and stones it usually signifies that it is
to be left for good and there is no one to carry
the bones about so that the bones will not be
removed. There is no belief as to any ill result
to the spirit from this practice. Usually, how-
ever, the bones are expected to be dug up.
In this case the body is covered by a sheet of
paperbark, the earth replaced and poles and
stones placed on top to stop dingoes, lizards
or dogs eating the flesh. If a camp dog does eat
the flesh it becomes wukundi, and may be
killed. If it is not killed it will be put through
a cleansing ceremony and any food that the
dog catches before the cleansing can only be
eaten by a male owner of the dog. Even after

the restriction has been removed the wife of
the owner or other women will have to make
a ceremonial presentation of food to the marra-
morkoimirri as soon as she eats food the dog
has killed.

A grave may be located in one of three
places. Where the deceased is a child, it is
often buried in the camp of the parents who
sleep beside the grave until exhumation. If it
is an adult the body is usually buried outside
the camp, but it may may be in it if the people
plan to abandon camp immediately.

In the southeastern part of the Murngin area
around Blue Mud Bay there is the third kind
of location: the collective burial ground.

Case 2. Visit to communal burial ground. On
October 18, 1935, Thomson visited a communal
burial ground or wukundi place at Blue Mud Bay.
It was situated on the edge of Marrakulo territory
at a place called Mange'yall, 5 km from the
Aborigines' camp. [Warner (1958: 49) refers to
the people of this area as Marungun and notes
that their Vaterhole' and 'country' are called
Mangaia which he glosses as 'stench of a dead
man' .J

A Marmariny man had died some 2-3 weeks
earlier. His body had been taken to this place
and put on a platform. The platform stood about
1-50 m high with the body upwards and roughly
covered in paperbark. The ground around the
platform was sculpted into a wandjur pattern. The
body was sharing the platform with a number of
other bones from skeletons, most of which were
covered with red ochre. In this case the head of the
body was placed towards the east so that the malli
[shade or shadow] could go to Buralko [the land
of the dead].

On this day Thomson visited the burial ground
in the company of five men: Raiwalla, his Mild-
jingi companion; Taudauongo, the actual elder
brother of the deceased; an old Ritarango man;
Djimbaron, a Dai'i man; and Marrilyanwi, a
Marmariny clansman who called the deceased son
and had been one of the kong wukundi. The
other kong wukundi were Liawulpul a Bidingal
man who called the deceased du'wai (FZS) and
Marakuri a korrong of the deceased (FZDS).

The party set out from the camp travelling
across salt pans. A line of fires was burning in
the short grass between the camp and the burial
ground. As the grass was sparse and short and
provided neither food nor cover for the animals
this was surprising. On enquiry it was found that
the fires had been lit to 'block in wukundi —
might be smell go all round', i.e. to cut it off from
the camp to which it is believed to be more or
less connected by smell. When the party was
within 0*50 km of the place the Aborigines re-
quested Thomson to leave the water he was
carrying lest the malli [shade] of the wurkaidi
(larvae ancestor) might 'go into it and make

100

NICOLAS PETERSON

(him) sick (rerri) . . .'. They translated the term
rerri, generally used for sickness of any kind, as
leprosy.

None of the Aborigines carried spears or spear-
throwers which was most unusual; this was said
to be one way of helping to avoid sickness. As
they neared the spot, Marrilyanwi took charge.
Thomson was told not to stay too long or go
too close lest he should fall sick. They all ap-
proached the platform by a roundabout path to
take them upwind. They conversed in whispers
and walked slowly with the arms folded to avoid
being 'flash'. The little outcrops of stone that
appeared on the flat salt pans were carefully
avoided. A couple of hundred yards off they halted
and Marrilyanwi rubbed his hands under his
armpits and then over Raiwalla's arms and legs
before kneeling down and biting his knees and
shins all the way down. Marrilyanwi then spat or
hissed in the direction of the wukundi place. He
did the same to Thomson. When they moved off
Raiwalla was told to walk behind Marrilyanwi in
his footsteps. When they reached the platform they
looked briefly and then moved away and turned
their backs while Marrilyanwi approached the plat-
form and smashed a pipe close to it in order to
pacify the malli so that it would not follow them.
'Chuckit smoke along him, that's all', explained
Marrilyanwi. They left and the Aborigines washed
in a salt pan a few hundred yards off. 'Wash'em
sweat, ground, that maggot him bite you and me—
malli — you and me no been see that malli\ one of
them commented.

On October 24, Thomson visited the burial
ground again with a Ritarango man, Wuruwul,
who had not been there before. Raiwalla and
Thomson did not have to go through any of the
procedures of the previous visit, although they did
approach upwind again. Wuruwul, a stranger to
the place, was very fearful of sickness, particu-
larly because he was somewhat fat. As a precaution
against sickness he had his knees and elbows bitten
(see Plate 5) and left the area well before the
rest of the party.

The alternative to ground burial is exposure
on a platform, either built in a tree and called
djamba or free standing like the kind used in
house building and called katauwurro. [Warner
states (1958: 433) that the body on a platform
is placed face up so that when the adominal
wall breaks the intestines will not fall.] Thomson
found that although this was true for the eastern
half of the area the people to the west of the
Ritarango place the body on its front.

The choice of burial mode depended on
several factors. Small children and old people
were usually put in the ground and active people
in their prime, male or female, were placed
on platforms where the flesh dries more quickly
and the bones become cleaner. If a person were
killed in a miringo raid they were usually put

in a tree by their relatives so that the people
could leave the area immediately.

There is a third mode of disposal found
among the Burara who Thomson reports as
eating young men, women and children, after
roasting in ashes [although Thomson did visit
the Burara it is not clear how much of the
following information was obtained by talking
with and observing Burara people and how
much was supplied by their Glyde River neigh-
bours who hold the Burara in low esteem].

The dead are eaten by all relatives with the
exception of M and MB because the 'two fella
been carrim along bindji [belly]'. Bodies to be
eaten have an incision made in their left side
through which the viscera are removed. The
liver is eaten but the heart, penis and vulva
are dried and carried in a special small basket
(pulupur in Burara) or in a matjitji to increase
hunting effectiveness. The lungs and stomach
are buried.

Because the body is eaten the western Burara
do not paint it, although the eastern Burara,
under the influence of their western neighbours,
the Wallamango and Yarnango, do. These latter
groups rarely eat their dead but do inspect the
internal organs, for signs of sorcery such as
sores (tjitjt) on the kidney, heart or liver, by
making an incision between the crest of the
ilium and the last rib on the left side. Now that
there are marngit medicine men in the area —
a new tradition from the south [see Thomson
1961] — the people no longer perform this kind
of investigation.

POSSESSIONS

The Djinang speaking peoples pull down the
deceased's hut and eventually set fire to it when
the bones have been exhumed. The main pos-
sessions of a person, such as his spears or a
canoe he has made, are treated throughout the
area in the same way as people. They are sym-
bolically cleansed at a mantjarr ceremony and
in the case of canoes rubbed with red ochre
often on two separate occasions. If a man has
been speared, his possessions are broken and
pieces given to his relatives in camp which an
informant interpreted as 'that mean I push all
these people go for fight'. Each piece of broken
possession used in this way is called maidjaballa.

MEM. NAT. MUS. VICT. 37 PLATE 5

Wuruwul having his elbow bitten as a precaution
against sickness prior to visiting the communal
burial ground in Blue Mud Bay (Case 2).

MORTUARY CUSTOMS

101

If a person does not want to fight he makes
a new dilly bag and gives it back to the man
(usually the elder brother of the deceased),
who distributed the maidjaballa. The broken
pieces are given most frequently to relations
in the categories MF/FMB, MMB, FZS,
MBS, ZS.

If a person dies from a cause other than
spearing his possessions are also broken up and
distributed to various other residential groups
where there are close relations of the deceased.
These possessions are then used in the clean-
sing ceremonies.

Thomson reports a case where following the
death of an important man a restriction was
placed over a large area of land.

Case 3. An area of land placed under restriction.
While at Katji [on the mainland south of Milin-
gimbi] camp in January 1937 Thomson noticed
red blazes on the trees along the path to Derby
Creek. These were to free the area about the Katji
River from a restriction that had been imposed
at the death of an important man.

The restriction had been imposed at the death
of Raiwalla's father-in-law [Raiwalla, a Mildjingi
clansman, was Thomson's guide and friend] be-
cause of his influence when alive and because
he had spent much of his life in that area. The
restriction was not removed until after the bukubut
[exhumation ceremony], when the trees were
painted and a lire lit to burn off the grass and
cleanse the area- — both literally and figuratively.
The fire was started by burning his old camp
with a fire of ironwood. After the burning off the
women went out and collected root foods in the
area and brought them back to the son and bro-
thers of the deceased. If any outsider eats food
from the area while the country is under the
restriction the close relatives resent this and try
to kill the offender by sorcery or with an actual
war party (miringo). Such restrictions do not apply
in remote areas but only when important people
{yarkomirri) die in the vicinity of an important
camping place. The death of the wife or daughter
of an influential man can also lead to the same
restriction Tjambarapoingo, Kopapoingo and
Ritarango peoples have the same custom.
CLEANSING CEREMONIES AND GROUNDS
Cleansing ceremonies of the kind described
in Case 1 are held at several stages following
death. After burial the clan song cycle of the
deceased is sung about a circular sand sculpture
and all men, women and children present,
together with the larger possessions are dusted
with heated leaves (mantjarr) to drive the nialli
(shade) of the deceased away, to render the
hunting weapons effective and the other pos-

sessions safe to handle.

At a second ceremony of similar form the
participants throw pieces of the deceased's
possessions into the fire burning in the small
depression which forms the focus of all the
cleansing ceremonies at this stage. At this
second stage the women usually dance while
the men are singing the clan cycle.

A week or two later the third cleansing
ceremony, called bukulup (forehead-washing),
is held. The small circular sand sculpture used
in the previous ceremonies now gives way to
a much larger and more elaborate representa-
tion of the clan well. Starting in the early hours
of the morning the men sing the clan songs
and then once the sun is up the close relatives,
male and female and the kong wukundi, wash
standing in the well and rub red ochre over
themselves. This ritual frees the kong wukundi
from all tabus. The patriclansmen make a pre-
sentation of food to members of the opposite
moiety.

The sand sculptures are also used in two
other contexts. Most frequently in the curing
of sores or wounds but also around graves and
burial platforms (see Case 2). Several clans
may share the use of a particular design and
a single clan may have several designs relating
to different places with differing degrees of
importance. Figure 1 shows sketches of six
grounds seen in use by Thomson.

EXHUMATION AND FLESH DISPOSAL
After a month or two the bones are exhumed.
The men who perform this task are also known
as kong wukundi [possibly also as kong djok]
but are not necessarily the same people who
buried the body. They may be of either moiety,
with the reservation that a man may not assist
in the exhumation of his siblings.

The grave is usually dug out either with bare
hands or with a sharpened stick. Among the
Djinang speaking groups only males are present
at the grave. One will sing and another play
the digeridoo. Among the people to the east,
women are present at the exhumation and dance
while the men sing.

The treatment of the flesh varies with the
kind of burial and the area. Among the peoples

102

NICOLAS PETERSON

SOME WANDJUR SAND SCULPTURE DESIGNS

(Running from left to right and top to bottom)

1. Marango clan (dua moiety) wandjur representing
the bee hive yarrpain [referred to as 'long-nose
sugar bag' in Aboriginal English on account of the
relatively long entrance tunnel]. The central
circle represents the eye of the clan well; the
rectangle surrounding the well and the area im-
mediately above it is referred to by the sacred
name bambula. The small rectangle had a pole
483 mm tall erected in it, called warrinman, repre-
senting an ancestral hero. The ground was used
on July 24, 1937, at Milingimbi for the cleansing
of two small girls. The same design may also
be used by Tjambarapoingo speaking clans.

2. Birkilla clan (yiritja moiety) wandjur representing
the bee hive birkurda at a place called Yarrakka
in Arnhem Bay. The small circle at the end rep-
resents the entrance (ngorro — nose) of the hive.
[This wandjur should be compared with the illustra-
tion published by Thomson in the 'Illustrated
London News' for February 25, 1939, page 294,]
This ground is the most elaborate form of the
wandjur and only used for important men; others
have a simplified, but recognizably similar, ver-
sion. The ground was used on August 7, 1937,
to cure a Birkilla/Kopapoingo man with a sore.

3. Birkilli clan {yiritja moiety) wandjur representing
waitjura [? a fish]. The piles of white sand are
sores (tjitji or mapan) made by a crab (mirriya
or katjirri) . The location referred to is Karraparra
in Blue Mud Bay and the design is also used

by Yituwa clansmen of that area. The ground was
used on August 7, 1937.

4. A dua moiety wandjur used by Liagauwumirri.
Maiyarrmaiyarr and other Tjambarapoingo speak-
ing groups united by the track of the Djanggauwo
sisters. The ground represents springs left by the
sisters whether they thrust the 'yam' sticks into the
ground. The springs are called milmindjarrk [and
are marked by being freshwater sources in areas
of salt surface water. The arrows appear to indi-
cate the direction of flow of the waters beneath
the wells]. The ground was used on July 30-31,
1937, at Milingimbi for a Maiyarrmaiyarr man
who was drowned when a canoe turned over
during a storm in the Cadell Straits. There were
six people in the canoe: two men escaped but
a blind man and a second man [it is not clear
which was the Maiyarrmaiyarr man] with his son
and daughter were drowned because they were
encumbered with turtle hunting gear. The ground
measured 15 m overall.

5. Kolumalla clan (dua moiety) wandjur represent-
ing mar'ndi [?] The ground was outlined in white
sand and used on August 14th, 1937.

6. Warramirri clan (yiritja moiety) wandjur repre-
senting a whale, woimirri. The small circle is the
rectum above the tail. The central rectangle is both
the whale's stomach and the manotji or eye of
of the clan well. The soil forming the outline was
raised up 100-125 mm and whitened with sand.
This ground was used on August 14 and 15,
1937, to cure a child of the Wunguri clan who
had sores. The child's full MMB came from the
Warramirri clan.

MORTUARY CUSTOMS

103

east of the Glyde River, including the Kanal-
pingo and Djinba, flesh from a grave burial
is put back into the ground. Flesh from a plat-
form burial is placed in paperbark and left in
a forked tree nearby to be destroyed naturally.
The platform itself is pulled down and buried
in the sand sculpture in which it was standing.
Among the Djinang, Mildjingi, Balambi,
Wullaki, Burara and all groups westward of
the Glyde, the flesh is kept and at the bukubut
ceremony placed in a special hollow log called
larkan djammurmur.

Case 4. Larkan djammurmur form of flesh dis-
posal. Early in the morning of November 11, 1936,
the Wullaki group at Katji started to sing in
preparation for a bukubut. The Wullaki people
were joined by some Milli'ereng clansmen because
the dead woman's mother was of this clan.

The body had been buried in the ground. A few
days earlier it had been exhumed and the flesh
roughly stripped from the body and wrapped in a
paperbark bundle. The bones were in a second
bundle. The men had then cut a tree for the
larkan [coffin] and the women collected cycad
nuts for a food presentation. While the cycad
nuts were being leached the larkan was fashioned
and a marraidjirri meri (an effigy of an ancestral
spirit) made. The preparation of the larkan in-
cluded singeing it, cutting the spikes on one end,
and painting it. On the morning of November 11,
1936, the men were seated in the shade of a
clump of trees some distance from the camp with
the larkan. The bundle of flesh was apparently
[this not unequivocally clear in the notes] already
inside the coffin. The wrapped bones and a mar-
raidjirri meri called Kanangalkngalk were also
nearby. This ancestral spirit was responsible for
the people using a log coffin.

An informant explained that in the distant past
(millegidji) there were spirits (meri or morkoi)
that were neither animals nor men and who still
live in the bush today. These spirits were never
men but a race of their own. One of these spirits,
Kanangalkngalk, is still alive today and some Wul-
laki people even claim to have seen him in the
monsoon forest. Kanangalkngalk has two wives
and some children, none of whom is a threat to
the people like the spirits of deceased human
beings. The marraidjirri meri represent Kanan-
galkngalk.

They say that in the distant past Kanangalkngalk
cut down a hollow tree. The tree fell and as it
fell water started to pour out of it. He tried to
hold onto the log as the water flowed out but his
fingers slipped and the log moved off like a fish.
The log cut the ground as it went allowing the
water to flow. Along the way the log heard a
buralla [publicly used bull-roarer) sing out. Then,
perhaps because the water told it to, the log
went underground at Katji carrying earth and
water with it as it went. The log wanted to go
down towards the sea but found the ground too
hard so it came back and let the water go. From
the end of the Katji lagoons, where

he turned back, there is no deep water but only
transient flood waters at the end of the wet season.
The log came back to Katji and decided to stay
where the deep pool is beside the camp. At that
place he gave himself a name: the log said, 1 am
djammurmur larkan." The people today reflect on
the fact that they take fish, water snakes {Hypsir-
hina) and wild taro from Katji where the larkan
walked around. That is why they paint them on
the log coffin and cut the long 'fingers' into the
mouth of the log, representing the jagged end
which resulted from it breaking the ground and
making Katji River.

The men around the larkan began to sing a
song about the wullawarri fish drawn on the log.
Then they picked up the log and danced with it,
making short lunges and rushes, replacing it on
the ground at the end of each movement. (See
Plate 6.) The log was then erected on the open
ground near the camp and the marraidjirri and
bones were carried toward the camp by two old
men. The dance that followed was called after the
spirit, Kanangalkngalk meri. The main body of
men danced forwards looking for the meri carried
by an old man, who represented the male spirit.
The one who carried the bones represented a
female morkoi [spirit] [Kanagalkngalk's wife?].
The first old man kept dancing forward with the
meri to reassure himself that the other man had
the bones.

The women danced their slight shuffling dance
from one foot to another on the fringe of the
dance area. The dance concluded with an old man,
dalkargrining [dalkarramirri] calling the big names
of the maraiin of the deceased and of her country.
Then the bones were handed over to the actual
younger sister of the woman's mother, the real
mother having died. As she received the bones in
their bundle the marraidjirri meri was placed on
top and she walked off with the whole lot.

At a later date the string on the marraidjirri
meri is removed and made into ngaimbak [arm
bands] and used in decorating a bati giwillir [a
kind of men's basket]. This basket is then pre-
sented to the man who made the string. The con-
clusion to the bukubut comes sometime later. A
presentation of food is made to the kong wukundi
by the close relatives of the deceased, usually the
F. duwe (either FZS or D), MB and EB if it is
a man that has died but not in the case of a
woman.

This food is not eaten bv the full F or MB
but FEB and FYB, duwe and kalli (MBS/D) do
share in it.

In the past the Djinang people removed the
flesh from the buttocks, washing it free of the
soft and more putrid surrounding flesh and tied
it up in paperbark. Later these parcels of flesh
were cooked outside the camp, wrapped in
grass and paperbark and hung around the neck
to increase hunting effectiveness. Some people
would go a step further and soak the flesh in
a mixture of honey and water and then nibble
a fraction with their eyes closed.

104

NICOLAS PETERSON

The bones are washed and wrapped in paper-
bark, of if the flesh is not entirely removed,
left exposed in a forked tree. During the 1-2
weeks before the bukabut ceremony the bones
remain outside the camp and may not be seen
by the women.

The kong wukundi camp apart for several
days. The wetter the body the longer the period
of restriction. If it is particularly sloppy the
men eat with a bone point (pringal) or any
sharpened stick because the fluids will have
penetrated their finger nails making them smell
for some time.

A day or two after the exhumation the kong
wukundi participate in a cleansing ceremony
singing all night and washing in the morning in
a sand sculpture. After washing they cover
themselves in red ochre. Several days later they
have a clan design painted on them which re-
leases them from all restrictions.

Case 5. Exhumation. On January 13, 1937, the
bones of a Djinang man named Lamieri, dua
moiety, buried at Gillere in Millierieng territory
were exhumed fsee Plate 7).

Only a few people went to the area of the grave
where the man had been buried 6-7 weeks before.
Those not directly involved in exhumation stood
upwind of the grave.

Two men were involved in handling the bones.
The man who removed the bones was the adopted
father of the deceased from the same country
[i.e. clan] named Balambarri. He was assisted bv
Makani a Mildjingi man a 'ZS 1 of the deceased,
who was married to two of his daughters.

On the way to the grave there was some discus-
sion as to whether the body was soft enough
yet to make the removal of the bones easy. The
grave itself was unmarked except for a plain cir-
cular sand sculpture near the head of the grave
and a heap of wood lying on top to keep the dogs
off. The body was about 1 m below the surface,
lying face down on a layer of grass and completely
extended. The grave itself was in loose, well-
drained sandy soil about 90 m from a creek. The
soil was removed largely with bare hands but use
was made of a canoe paddle that happened to have
been in the camp and brought along.

An old clansman of the deceased, a classi'ficatory
F, sang_ to the accompaniment of clapsticks. In
Kopapoingo and Tjambarapoingo ceremonies
women are present and dance during the exhuma-
tion but not among the Djinang. The kong wu-
kundi examined the body to see that the flesh
was sufficiently decomposed and finding it was, the
assistant, Makani, went off to get some water in
a paperbark trough.

The deceased had been an old man of not much
standing so he had not had a clan design painted
on his chest.

The adopted father removed the bones, starting

from the feet and working upwards. Makani poured
water over the bones as the first man washed them
thus reducing the period he would be wukundi
[tabu]. The head was picked up last and washed
by pouring water in through the foramen magnum.
Each bone as it was picked up was placed on a
sheet of bark beside the grave. When they had all
been removed the grave was filled in again. The
two men washed and smeared themselves with
white paint from head to foot: 'everybody no
more want to smell*. Red ochre is only used ixi the
final cleansing. During the night a sand sculpture
1-60 m in diameter was made at Makanrs camp
and a ceremonial washing called bukulup carried
out on the following morning (see Plate 8). About
sunrise the two kong wukundi washed by pouring
water over one another's bodies and then smearing
red ochre all over themselves, their spears and
spear-throwers and immediate possessions. This
freed the men and their weapons from tabu.

It is usual to wait two or three days before
holding this ceremony, but as the camp was break-
ing up on the following day it was completed
straight away.

MARRAIDJIRRI MESSAGE STRINGS
Marraidjirri is the general term for a class
of decorated strings whose most common use
is in the mustering of people for exhumation
and final disposal ceremonies (see Plate 9).
Marraidjirri strings differ from mamgarai
strings in that although many of them incor-
porate hair of a dead person they are largely
made of fibre string and are used in a different
way, (for marngarai strings see under 'Hair'
above).

Each clan has its own marraidjirri forms
representing totems associated with the clan
(see Table 1). Generally there are several
forms of the string-like marraidjirri which are
classed together as bogongo and spoken of as
'small' in contrast to the elaborate figure em-
blems such as that mentioned in Case 4 which
are referred to as big (yindi).

Besides being used to gather people for
mortuary ceremonies they are also associa-
ted with circumcision ceremonies, the social
development of children and love magic.
In circumcision ceremonies they are used to
gather people. The second usage result from
the first occasion on which a small child
picks up any natural object such as grass,
a shell, fruit or small lizard and gives it to
to its parents. This object is then tied into a
small bundle and sent off to an acquaintance
both geographically and socially distant. This

MEM. NAT. MUS. VICT. 37 PLATE 6

«-* * 4f"*- ■***

Wullaki men taking the larkan coffin to the camp
for erection (Case 4).

MEM. NAT. MUS. VICT, 37 PLATE 7

The exhumation of a Djinang man (Case 5).

MEM. NAT. MUS. VICT. 37 PLATE 8

Ritual washing in a wandjur sand sculpture of
two of the men who participated in the exhuma-
tion shown in plate 4 (Case 5).

MEM. NAT. MUS. VICT. 37 PLATE 9

On August 20, 1935, this messenger, a man of
the yiritja moiety arrived at Milingimbi wearing
a dua moiety marraidjirri message string. He had

come to call the people to Elcho Island for the
final disposal of the bones of a Tjamborapoingo
man.

MORTUARY CUSTOMS

105

person fashions the object into his own clan's
large marraidjirri and presents it to the child's
parents in a large public ceremony. The parents
then make a payment of traditional wealth of
considerable proportions to the maker of the
marraidjirri. On completion of the ceremony
the string is removed from the core about
which it is bound and made into arm bands or
used for adorning certain kinds of men's baskets
( bati mindjalpoi ) . Some marraidjirri strings
may be used in love magic after they have been
used in one of the foregoing ceremonies A
sweetheart or errant wife is believed to be
impelled to follow the man involved when the
marraidjirri string is looped over her hands.

The strings sent out to muster people for
ceremonies are really representations of big (or
proper) marraidjirri constructed around wooden
or sometimes paperbark centres and used at
the bukubut (see Case 4). These solid emblems
look like rangga (the secret totemic emblems)
as an informant observed to Thomson, but may
be seen by women and children and are said
to represent the clan's ancestral morkoi or
spirit being. This morkoi is different from the
clan's ancestral hero, wangar. However, as with
the clan totem, the string covering is called
buy it and is equated with flesh; in its broadest
sense it just means covering. The core is re-
garded as maraiin (sacred) and identified with
the bones of the skeleton.

TABLE 1

Some of the small marraidjirri strings
(bogongo) used by various groups

CLAN/GROUP

Wanguri

Birkilli

Tjambarapoingo

Kanalpingo

Liagallauwumirri

Mandalpoi
Ritarango
Durrilli
Obulkarra

Mildjingi

NAME and DESCRIPTION

Yorko — a round root called Kalun
in Kopapoingo (Cissus carnosa)

Komulo — the great billed heron

Ku'ak — a small bird

Yukuwa — the root food Vigna
vexillata

Tjarrak — a tern {sterna sp.)

Kalliwur — a large white lily

Wititj— Snake

Ku'ak — a small bird
Malka — -bee (Trigona sp.)
Yorko — round root (Cissus

carnosa)
Kurungur — a small cloud and also

the wild bean Ipomea pes-caprae

BUKUBUT FOLLOWING EXHUMATION OF
BONES ONLY

When the food for the bukubut ceremony is
ready, a week or two after exhumatiom a classi-
factory ZS goes into the bush and picks up the
bones. Meanwhile the marramorkoimirri pre-
pare the ground, knoww by different names to
different clans:

Liagallauwumirri call it birlimbil

Djeranggoikoi „ „ djirrkurul (of

bulmantji or
shark)

Birkilli „ „ yallandu (from

Bukunda, a
place)

Mildjingi „ ,, manitji

When the 'ZS' carrying the bones, ap-
proaches the ground just outside the camp
where the ceremony is to be held, the men
begin to sing. The marramorkoimirri and a
great crowd of more distant kinsmen dance
around with spears poised and jab these at
the bones in their paperbark wrappings as
they lie in the sand sculpture. 'Him want spear
that one, open him — wangar (totemic ances-
tor) been do.' Not all clans carry out the
ceremony in this way. The western ones, Lia-
gallauwumirr, Mildjingi and Djinang only sing.

The Birkilli, Daigurgur, Ritarango and al-
lied groups customarily spear the parcel, and
do the same again when they are holding the
final disposal ceremonies. They open the paper-
bark with the spears and then sit down and
wash the bones before placing them in a new
wrapping, and hanging them from a forked
stick standing in the middle of the sand sculp-
ture. At this stage the women and children
may not see the bones although they can later
on when they have been red ochred.

From the late afternoon onwards through
the whole night the men sing and complete the
bukubut ground where the bones will be pre-
sented to the woman who is to carry them.
This is usually the actual FZ, EZ, adult D, or
mother if the deceased is a young child. If
the bones of an older person are given to a
mother it is always to a classificatory mother;
they are never in the custody of a sister

106

NICOLAS PETERSON

although she may handle and carry them. In
the morning these relatives and the MM will
dance near the song group which is usually
composed of EB, YB, F. The bones are then
handed to the MF/FMB, ZH or MMB. Most
commonly, it is to the ZH who, then hands
the bones to the deceased's FZ who will carry
them during the following weeks.

Previously the F, S, ZDS and ZS of the
deceased will have made a bark container,
tarra, decorated with the deceased's clan design
for the bones to be carried in. The Mildjingi
and Liagallauwumirri do [may?] not have this
custom. There is then a ceremonial present-
ation of food by all the helpers in the various
stages of the ceremony to the marramorkoi-
mirri.

Often the skull is not placed in with the
other bones but carried separately by WB or
another ZH. The bones are carried for 1-5
weeks. At the end of this period they are again
hung from a forked stick in camp and only
moved on shifting camp, until the final disposal
of the bones in a hollow log coffin ceremony.

Case 6. A Wullaki hukuhut ceremony. A bnkuhui
ceremony at which the bones of a dead Wullaki
speaking man of yiritja moiety were handed over
to the deceased's sisters was held at Katji on
October 3 and 4, 1936.

When Thomson arrived in camp late in the
afternoon the ceremony was about to begin. The
bones were hanging in a small shade (kurngan)
along with the marraidjirri meri. The marraidjirri
represented a wasp's nest called banal and was
decorated with a picture of the little green pigeon,
work'miringo (Chaleophaps chrysochlora). This
bird is associated with the paper wasp in areas
of monsoon forest.

Although the deceased man was Wullaki the
bukubut was carried out with a Mildjingi song
sequence because the Wullaki relatives had handed
the bones to the Mildjingi [Thomson has Raranggal
malla at this point in his notes, but on the first
page he equates Mildjingi malla with Raranggal
malla] clansmen as a friendly compliment.

The ceremony began with singing to the accom-
paniment of clapsticks and didgeridoo and con-
tinued for some hours into the night until every-
body was supposed to be asleep. Each man then
seized a torch of lighted paperbark and started to
dance, first encircling the shade with the bones
and marraidjirri in it, and then moving into the
camp crying berk! berk! berkberk ko ye'h ko yeh.

They encircled the whole camp where everyone
pretended to remain sleeping and then trotted back
in single file to the shade. This was the dance of
the flying fox. The singing then continued. When
they began the song about the jungle fowl (gulla-

uwurr) the men started to dig a long serpentine
path, which eventually measured 42 m, from the
shade to the point where the men constructed a
representation of the bird's nest. At intervals the
men working on the road and the nest cried out
kurrkun kurrkiin djue wurak — grr'rr in imitation
of the jungle fowl; these cries were heard inter-
mittently through the night. Jn making the nest
mound the men imitated the movements of the
bird by crouching low.

Early in the morning the marraidjirri was
brought out (see Plate 10). The first dance was the
wasp dance. While most of the men danced,
two of their number darted out towards a
man who held the marraidjirri at arm's length
on a spear-thrower and pretended that they were
being attacked at the wasp's nest. The men were
meant to be looking for yams in the monsoon
forest and to be driven back by the wasps. The
women danced at some distance with the common
rhythmic shuffle known as luku wankain 'ngorro.
When the dancers reached the jungle fowl nest at
the end of the path they called out the big names
associated with the deceased. These were both the
deceased's maraiin (sacred) names and those of
the Mildjingi clan. As the calling finished the
bones were handed over by the deceaseds WB
(Bulambirri) to the dead man's full sisters. [They
would not be the custodians of the bones].

During the period in which bones are carried
they are thought to indicate the approach both
of news bearers and of revenge parties. If the
bones make a light tap against the bark con-
tainer this indicates the approach of a person
with news of some kind. If the tap is loud it
announces the approach of a war party.

Once or twice before burial in the hollow
log coffin, the bones are taken out of the
tarra and red ochred and this may be associ-
ated with the change of tarra too. After the
second red ochring the mother may become
custodian of the bones. This painting with red
ochre removes wukundi from the woman who
carried the bones intially.

HOLLOW LOG COFFIN
The holding of the final ceremony is decided
in this way. The father or his brother, asks the
relatives carrying the tarra whether they are
ready to make the coffin. If they agree prepar-
ations are made for the ceremony. Frequently
the relatives carrying the tarra feel the need
to make an excuse to agree and say that they
are ready because they have carried the bones
for a long time without help from anybody
else. The coffin is made by F, EB, YB and

MEM. NAT. MUS. VICT. 37 PLATE 10

Dancing with a wasp emblem in a Wullaki buku-
but (Case 6).

MEM. NAT. MUS. VICT. 37 PLATE 11

A hollow log coffin ceremony in Arnhem Bay,

1937.

MORTUARY CUSTOMS

107

MMB, all members of the deceased's patri-
moiety.

Hollow log coffins collectively referred to
as dupan (hollow) differ in size, ranging from
1.25 to 4.50 m. and different clans call them
by different names (see Table 2).

TABLE 2

The names used by different groups for their
hollow log coffins

CLAN/GROUP NAME

Ritarango
Mandjikai Wurrwurr

Birkilli Djallumbo — associated with a wad-

ing bird found on the salt pans

Tjambarapoingo Daimirri — a hollow log thrown by
an ancestral hero into the sea
and transformed into a hollow
stone outside Buckingham Bay

Liagauwumirri Mululu

Liagallauwumirri Bardaru — Associated with the crow
and the milky way

Marango Kallangurr

Kanalpingo Larradjadja

Mildjingi

Warramirri Kapalla — associated with the fun-

nel of a steamer and the blow-
hole of a whale

Tjapu Larrakit

However, all are made in the same way. A
tree which has been hollowed out by termites,
is cut and cleaned by burning. A circle is in-
cised 300-600 mm from the top end and
two small holes cut diagonally opposite each
other 50-75 mm from the top. The circular in-
cision is called derong and always painted yel-
low on yiritja coffins and red on dua coffins.
The two holes are called eyes and serve to
commemorate the fact that the coffins w r ere
originally made by each clan's spirit ancestors
and in some way personify them.

After the coffin has been shaped it is moved
into a large shade where the men work on
painting it. As the painting nears completion
the people gather for the final ceremony. Each
evening there is singing and dancing. On the
final morning the bones are taken from the
tarra and covered with red ochre. The skull
is then painted with the clan design. The coffin
is brought out and placed at an angle with one
end supported on a forked stick. Inside a sand
sculpture of the clan well a close male relative
of the deceased, often a korrong (FZDS) or
moralkor (MMBS) breaks the long bones and

the skull before placing them in the log. Final-
ly the log is erected (see Plate 11).

The bones of several people of both moieties
may be placed in the same coffin. The coffins
are left standing, eventually decaying and dis-
appearing without trace.

References

Thomson, D. F„ 1961. Marrngitmirri and Kalka—

medicineman and sorcerer — in Arnhem Land.

Man 61: 97-102.
Warner, L., 1958. A black civilization: a social study

of an Australian tribe. New York: Harper and

Brothers.

Notes

1 The Donald Thomson Ethnographic Collection
was donated to the University of Melbourne by Mrs
Thomson following the death of her husband in May
1970. By agreement the University has lent the Collec-
tion to the National Museum of Victoria where it is
now housed.

The principal purpose in preparing this paper for
publication is to draw attention to the ethnographic
riches of the Collection. No compilation of notes
can do justice to the vision that informed the field-
work nor create the interpretive synthesis that Thom-
son had in mind. Inevitably the immense ethnographic
detail of the fieldnotes can now be appreciated only
by a series of scholars who will be able to breathe
interpretive life into different aspects of the many
but unintegrated details that characterize all fieldnotes.

This introduction to the Arnhem Land section of
the Collection has been built on Thomson's notes for
a paper on 'Kopapoingo Death and Mourning Rituals'.
Within the general framework of the notes I have
added descriptions from his fieldnotes as case studies.
The presentation of both the text and the cases has
been kept as close to Thomson's own wording as
possible, but some alteration has been unavoidable
in the process of converting fieldnotes to continuous
prose. Further the notes cover several years and
during that time Thomson's understanding of the
language and life underwent substantial changes. In
particular, his spelling of words in the local languages
altered, so I have standardized on the later forms. A
number of details in the cases and in particular those
associated with the wandjur sand sculptures, have had
to be omitted since the meaning was obscure and
there was no simple way of setting out the informa-
tion, some of it possibly deriving significance from its
location on the page and its position relative to other
notes. Undoubtedly a scholar with particular know-
ledge of some of the clans' religious life would be
able to make sense of some of the notes that have
been omitted. For this reason any person working
intensively on a paricular aspect of the mortuary
customs described here or on the details of symbolism
in the life of a paricular group will have to consult
the notes themselves where they will find the odd
word or phrase that may be of significance to them.
I have enclosed substantive additions to the text by
myself in square brackets.

108

NICOLAS PETERSON

I received permission to prepare the paper for
publication from Mrs Thomson while organizing the
cataloguing of the ethnographic collection on a grant
from the Australian Institute of Aboriginal Studies.
Work on the preparation of this paper has been made
possible by an appointment as senior Associate in
Aboriginal and Oceanic Ethnology in the Department
of History in the Faculty of Arts of the University
of Melbourne. I am most grateful for the help I
have received from Mrs Thomson, Margret Darragh,

Gregory Dening, Alan West and Nancy Williams.
Special thanks are due to Judith Wiseman for her
unflagging assistance in all things connected with the
Collection.

2 It is uncertain whether this usage of wukundi is
correct. The primary reference is to places associated
with death that are tabu in some way (see Case 2).
Thomson appears to have extended the meaning to
cover other tabus associated with death.

*kmat

SRM

■

8H5

»?3SE|